STORKS, IBISES

and SPOONBILLS
of the World
This page intentionally left blank
STORKS, IBISES
and SPOONBILLS
of the World
JAMES A. HANCOCK
JAMES A. KUSHLAN
and M. PHILIP KAHL

Illustrated by
ALAN HARRIS and DAVID QUINN

Published with the support of the Brehm Foundation

ACADEMIC PRESS
Harcourt Brace Jovanovich, Publishers
London * San Diego * New York * Boston * Sydney * Tokyo * Toronto
First published 1992 by Academic Press.

Digital editions published 2011 by Christopher Helm Publishers, an imprint of A&C Black Publishers Ltd, 36 Soho Square, London
W1D3QY

Copyright © 2001 text Academic Press Limited

Copyright © 2001 illustrations Alan Harris and David Quinn

ISBN (epub) 978-1-4081-3500-6

ISBN (e-pdf) 978-1-4081-3499-3

All rights reserved. No part of this publication may be reproduced or used in any form or by any means - photographic, electronic
or mechanical, including photocopying, recording, taping or information storage or retrieval systems — without permission of the
publishers.

Visit www.acblack.com/naturalhistory to find out more about our authors and their books. You will find extracts,
author interviews and our blog, and you can sign up for newsletters to be the first to hear about our
latest releases and special offers.
 Contents
INTRODUCTION 1
CLASSIFICATION 5
CONSERVATION 15
C O U R T S H I P AND R E P R O D U C T I O N 20
F E E D I N G B E H A V I O U R AND ECOLOGY 29

SPECIESD E S C R I P T I O N S
American Wood Stork (Mycteria americana) 35
Milky Stork (Mycteria cinered) 43
Yellowbilled Stork (Mycteria ibis] 49
Painted Stork (Mycteria leucocephala] 53
Asian Openbill Stork (Anastomus oscitans) 59
African Openbill Stork (Anastomus lamelligerus) 63
Black Stork (Ciconia nigra) 69
Abdim's Stork (Ciconia abdimii) 75
Woollynecked Stork (Ciconia episcopus) 81
Storm's Stork (Ciconia stormi) 87
Maguari Stork (Ciconia maguari) 91
White Stork (Ciconia ciconia) 97
Oriental White Stork (Ciconia boyciana) 103
Blacknecked Stork (Ephippiorhynchus asiaticus) 109
Saddlebill Stork (Ephippiorhynchus senegalensis) 115
Jabiru Stork (Jabiru mycteria) 119
Lesser Adjutant Stork (Leptoptilos javanicus) 125
Greater Adjutant Stork (Leptoptilos dubius) 129
Marabou Stork (Leptoptilos crumenifems) 133
Shoebill (Balaeniceps rex) 139
American White Ibis (Eudocimus ruber) 147
Barefaced Ibis (Phimosus infuscatus) 157
Glossy Ibis (Plegadis falcinellus) 163
Whitefaced Ibis (Plegadis chihi) 169
Puna Ibis (Plegadis ridgwayi) 173
Sharptailed Ibis (Cercibis oxycerca) 111
Plumbeous Ibis (Harpiprion caerulescens) 179
Contents

Buffnecked Ibis (Theristicus caudatus) 183

Green Ibis (Mesembrinibis cayennensis) 189
Hadada Ibis (Bostrychia hagedasti) 193
Wattled Ibis (Bostrychia carunculatd) 197
Olive Ibis (Bostrychia olivacea) 199
Spotbreasted Ibis (Bostrychia rard) 203
Madagascar Crested Ibis (Lophotibis cristatd) 207
Sacred Ibis (Threskiornis aethiopicus] 211
Oriental White Ibis (Threskiornis melanocephalus} 217
Australian White Ibis (Threskiornis moluccd) 221
Strawnecked Ibis (Threskiornis spinicollis] 227
Waldrapp Ibis (Geronticus eremitd) 231
Bald Ibis (Geronticus calvus) 237
Black Ibis (Pseudibis papillosa) 241
Giant Ibis (Thaumatibis giganted) 245
Oriental Crested Ibis (Nipponia nippon) 249
Eurasian Spoonbill (Platalea leucorodid) 253
Royal Spoonbill (Platalea regia) 259
Blackfaced Spoonbill (Platalea minor) 263
African Spoonbill (Platalea alba) 267
Yellowbilled Spoonbill (Plataleaflavipes] 271
Roseate Spoonbill (Platalea ajajd) 275

APPENDIX 283
BIBLIOGRAPHY 329
INDEX 383
 Introduction

T
I here is no one word or concept that can describe fairly the diverse group of birds
numbered among the storks, the ibises and the spoonbills. Compared to the majority of
birds, they are among the larger and all have relatively long legs, necks and bills as
adaptations to their fundamental way of life—walking about in shallow water. However,
storks, ibises and spoonbills range in appearance from the elegant to the ungainly, in size
from the moderate to the huge, and in behaviour from the flamboyant to the secretive. Their large bills
may be massive or thin, straight or curved, tubular or flattened. Some species are found in large
spectacular assemblies and others in crepuscular isolation. Some of the species were once widespread
and common, but are now at or near to extinction. Others have always been rare, with limited ranges
and highly specialized ecological niches about which little is known. Each has its own mysteries yet to
be revealed, and each has its own claim on our fascination.
We have approached the task of preparing this book with some trepidation, not only because of its
magnitude but also because of the growing certainty that, unlike most heron species, an unacceptable
number of storks, ibises and spoonbills are facing extinction. Realization amongst the general public
that much of our avian population is becoming dangerously depleted in many, if not all, parts of the
world, has been slow to crystallize. Even amongst ornithologists, the true, and in many cases, horrifying
situation of these species is only now becoming apparent as information pours in from remote portions
of the globe. Information on rare storks, ibises and spoonbills is arriving at a rapid rate, and much of it
is not encouraging.
We have attempted to include as much of this up-to-date information on the changing status of the
species as possible. But again and again in preparing this book, we find ourselves emphasizing how
these birds are in competition with humans for space and resources. With the world's human
population expanding, the space left for large waterbirds has shrunk so rapidly in so few years that we
have barely yet come to grips with the devastation that has been wrought. Two particular scenarios
tend to repeat themselves among the species accounts. One is that of shy elusive species living as far
away from humans as they can in areas where they have had, until recent years, little contact with
people, but where they now find little habitat left. The other is of species which have lived in close
proximity to humans and have very often become dependent upon them for their very survival. An
example of the first scenario is Storm's Stork, about which we have learnt very little but which, because
of deforestation, might well have disappeared before we had fully realized that it existed at all. The
second group is epitomized by the Waldrapp Ibis which has been the recipient of every type of
persecution which humans are capable of inflicting.
Today we have reached a vital stage in the relationship between humans and large wading birds. The
task of conserving our environment and the birds within it can only be tackled if sufficient is known about
the needs of individual species within the increasingly hostile globe upon which we live. We hope that we
have managed to present as clear a picture as is possible of the precarious position of the various species of
storks, ibises and spoonbills around the world and, where possible, we have suggested appropriate action
to help their cause. A singular message from our review is: what is so desperately needed is protection and
management of the wetland habitats that are essential to most of the species we discuss.
We have endeavoured in this book to share with the reader our own fascination and concern for these
birds. We are fortunate to have been able to draw upon our own experience and that of our editors and
publisher to present what we hope is a readable and authentic account of most of what is known about
these species. We have been most fortunate, too, in obtaining the services of two highly talented and
dedicated artists, Alan Harris and David Quinn, who have provided portraits of each of the species
described. We hope this original artwork will contribute to both the enjoyment and usefulness of this
monograph.
2 Introduction
Figure 1. A Maguari Stork in full breeding plumage with
two nestlings approximately 2-3 weeks old (northeastern
Argentina).
In today's financial climate, meeting the expenses of
production and publication of a book of this nature, contain-
ing original plates, is thwart with difficulties. Vital to the
project was the support of the Brehm Fond and Mr Wolf W.
Brehm. We also are quick to acknowledge the seminal
contribution to the project made by Mr Charles Luthin,
working on behalf of the Brehm Fond, Vogelpark Walsrode,
and the Specialist Group on Storks, Ibises, and Spoonbills. A
volume such as this would have been impossible to publish at
other than an astronomical price without this generous
support.
We are fortunate to have been able to draw much of
our information and our insight from our own experience.
The authors have been privileged to travel extensively,
observing these birds on each of the continents they inhabit.
Between us, we have seen in the wild all but 8 of the 49
species we discuss. Two of us have spent many years study-
ing one of the sister groups of these birds, the herons
(Ardeidae), having previously summarized our knowledge in
a companion volume, The Herons Handbook (1984). Two of us
have spent many years studying the storks, spoonbills and
ibises themselves, the results of which have been published in
numerous papers addressed to our scientific colleagues; one
Figure 2. A Glossy Ibis near its nest (southeastern Aus-
tralia).
having spent over 30 years studying the storks and more
recently spoonbills and providing a fundamental perspective
on their behaviour, ecology and taxonomy, the other, over 20
years studying populations and communities of storks, ibises
and spoonbills on two continents, providing a comparative
perspective on their ecology, habitat and conservation.
To supplement our own observations we have consulted an
abundant although very uneven literature in an attempt to
synthesize what ornithologists before us have found about the
biology and conservation needs of these species. Much of this
literature is excellent; and, for present and future students,
we hope the extensive, yet selected, bibliography in this book
will be an entrance into this literature. The bibliography is a
collaborative effort, assembled by a team led by Dr Malcolm
Coulter (see below). One caveat: we would urge caution and
selectivity in the use of this literature, some of which goes
back many years. We have attempted to be selective and to
present the conflicting views, but undoubtedly will have
erred in our judgement in those cases where facts we
gleaned from the literature are really quite debatable.
We have also sought advice from a wide-ranging and
knowledgeable host of experts. The response to our requests
has been enormous, and we are deeply indebted to a very
 Introduction 3

large number of collaborators both amateur and pro-

fessional who have been so supportive, encouraging and
generous in sharing their insight and understandings.
Some of our trepidation is also due to our certain knowl-
edge that no matter how thorough our own efforts, the
literature, or our correspondence, a book such as this will be
incomplete and in some areas will be proved incorrect. Our
limitations can, and undoubtedly will, be quickly recog-
nized. The available knowledge of the biology of these birds
is highly erratic. Some species are very well known, whilst
others have hardly been studied at all. Our species accounts
reflect this discrepancy. Over and over again we bring to the
readers' attention significant gaps in our knowledge about
the most basic aspects of biology. It is the nature of such a
work to summarize, synthesize, pose hypotheses, and en-
courage further study and testing of these hypotheses. We
hope this book will inspire others to fill in the important
gaps in the knowledge and to test some of the hypotheses we
propose, so that the business of conservation can continue
more effectively. If the book accomplishes an increased
interest and study, we will be satisfied to have accomplished
our primary goaL
Range maps are always troublesome. We are conscious of
their summary value but have found, like many authors
before us, that illustrating the ill-known and changing dis- Figure 3. A Eurasian Spoonbill head-scratching (north-
tribution of many species is well nigh impossible. We have, ern India).
therefore, sought to show general ranges of each species,
reserving for the text a more detailed description. Even here
we have been handicapped by lack of precise information on presented will have brought together the sort of information
the range of a surprisingly large number of species, about which will provide encouragement and a structure for future
which little is recorded, and not least by the often unreliable researchers and observers.
recordings by collectors of the past whose poorly labelled It is our hope, too, that the emphasis we have placed on
specimens lack credibility. the dire need for conservation of many of these species, and the
Information on basic nesting behaviour is missing for wetland habitats on which they depend, will encourage res-
some species; and, for many, information on colour changes ponsible authorities to act on their behalf before it is too late.
during courtship are lacking or conflicting. Soft-part colours
of museum specimens are especially unreliable guides. In ACKNOWLEDGEMENTS
the field, information on courtship colours are seldom We are extremely grateful to all those who have, lent their
recorded in all phases of their development. Such knowledge assistance in the production of this book. As noted in the
as we have presented should be subjected to most careful Introduction, we especially acknowledge the support of Mr
further study by field-workers who are fortunate enough to Wolf W. Brehm, who has made it possible for original plates
come across them. Yet for species that change the colour of to be produced. We thank, too, our long-time colleague Mr
their skin during courtship, this information is vital. Charles Luthin, who nurtured the project, and because of
Voices and other noises are of importance to most of these whose efforts much of the information now available on the
species. Yet our understanding of their vocalizations is status of these species has been gathered by field-workers
primitive compared to that of other birds. However, in some throughout the world. We thank too the excellent artists,
cases the only reliable identification of a species that can be Messrs Alan Harris and David Quinn, for their outstanding
made is by voice. These are large birds, with mostly gruff contribution to the quality and usefulness of the book.
and non-melodic voices. We have rendered their calls in In actual production, we thank Dr Andrew Richford, our
English inventing such words as 'Cha-cha', or 'Hunk-hunk*. editor, Penny Robinson and Debra Kruse at Academic
These are recognizable by those who know them, but not Press for their considerable help. We also thank especially
necessarily by those who do not (in some cases, one simply Mrs Jane Ratliff and Mrs Bonnie Knight for their typing
had to be there). assistance and Mr Melvin Seid for bibliographic assistance.
With three authors combining their respective strengths We thank our colleagues who worked on the bibli-
and weaknesses, it is unfortunate that there has to be an ography. This was a collaboration lead by Dr Malcolm
order of authorship. Such an order does not imply the Coulter to assemble a bibliography for this group of species.
relative contribution of each. This was a true collaboration The present reference section is derived from the first edi-
of experiences and ideas. As an Anglo-American team, we tion of this bibliography, which is available on computer
express the hope that the individual and joint efforts here disk from Dr Coulter; the compilers of that bibliography are
4 Introduction
Figure 4. Strawnecked Ibis. Huge flocks fly north from southern Australia after breeding.
Malcolm Coulter, Koen Brouwer, Albert L. Bryan, Jr., M.
Philip Kahl, Catherine King, James A. Kushlan, Charles S.
Luthin, Marcel van Wieringen, and David P. Young, Jr.
We are very grateful to our many correspondents who
generously provided their unpublished information, obser-
vations, data and insight, and many of whom were kind
enough to review preliminary drafts of various chapters. We
take great pleasure in acknowledging their contributions:
Eduardo Aguilera, Bubphar Amget, J. Ash, George Archi-
bald, P.C. Bhattacharjee, William Belton, Bas Van Balen,
Hilde Bult, Navancet Bhatt, A. Brosset, Linda Birch, Keith
Bildstein, Walter Bock, Koen Brouwer, Benjamin Busto,
Nigel Collar, Miles Coverdale, Malcolm Coulter, Juan P.
Cueilo, Montserrat Carbonell, Finn Danielsen, J.H. Desai,
Peter Davey, Andrea Dinep, Catherine Deulofeu, Jon
Fjeldsa, Peter Frederick, Mercedes Foster, Paula Frohring,
Michael Gochfeld, Juan Carlos Guix, Alfredo Guillet, D.A.
Holmes, U. Hirsch, David Holyoak, Peter Hubbell, Cathy
King, Richard Klein, P. Kennerley, Kim Lowe, O. Lan-
grand, Charles Luthin, Catrina Martin and the Texas Parks
and Wildlife and Texas Colonial Waterbird Survey for the
use of survey data, Nigel Mathews, Burt Monroe, David
Manry, D. Melville, Adam Mrugasiewicz, Gonzalo Morales,
Samuel Narosky, Eva Neumier, R. M. Naik, John Ogden,
Fabio Olmos, Storrs Olson, Raul Lopez Pacheco, Duncan
Parish, Kenneth Parkes, Jan Pinowski, D.
Pomeroy, E. Poorter, the late Peter Pugh, Crawford Pren-
tice, Ian Parker, Philip Round, Christina Ramo, James
Rodgers, Asad Rahmani, W.B. Robertson, Tom Roberts,
R.A. Ryder, Roberto Schlatter, Charles Sibley, the late
Helmut Sick, Marcel J. Silvius, Henrick Skov, W.R. Sieg-
fried, Sandy Sprunt, D. Scott, Holger Schulz, the late Ernst
Schiiz, Gary Stiles, P. Saikia, C. Sivasubramanian, S. Smir-
enski, Betsy Thomas, Hashim Tyalji, Daniel M. Taylor,
Tao Yu, Victor Terera, Emil Urban, AJ. Urfi, J. vanTets,
David Wells, Scott Wood, Marcel V. Wieringen, S.W. Win-
ter, M.D. Williams, Josef Witkowski, W.J.M. Verheugt.
We gratefully acknowledge the University of Mississippi
for its support for one of us (J.A.K.) and of this project, the
MacArthur Foundation for the award of a MacArthur Fel-
lowship to one of us (M.P.K.) allowing him to devote his
time to this project, and the staff of Cox and Kings Travel
Ltd for organizing travel arrangements in Asia for one of us
(J.A.H.).
NOTE ON RANGE MAPS
The following colour coding has been adopted. Areas where
the species concerned is a breeding resident are shown in
blue. Where a species is completely or partially migratory,
the breeding range is shown in red and the non-breeding
range in green. Arrows indicate principal migration routes
and ? areas where the status is doubtful.
 Classification

A S Y S T E M FOR C L A S S I F I C A T I O N
We are able to understand the world around us in part through our ability to classify objects and events
into manageable units and by establishing, in our mind at least, logical relationships among them. This
is especially true of the natural world; the classification of living beings is a specialized science called
Taxonomy or Systematics. As in our everyday efforts at pigeon-holing objects and ideas, the scientific
system of classifying organisms also has two goals, to identify and name the entities of the natural world
and also to describe the relationships among them. In developing a classification, taxonomists seek to
place together organisms that are related through descent from a common ancestor. Such organisms
share a common genetic inheritance that we expect to be expressed in various characteristics that we
can see or measure. Of course, characteristics may in fact, be similar without being derived from genes
shared with a common ancestor, and, conversely, there is no reason to suspect that all genetic similarity
should be expressed in characteristics we can easily observe or measure. An increasing ability to discern
biochemical differences, including characteristics of the DNA itself, provides additional measures of
genetic similarity.
The basic unit of taxonomy for organisms such as birds is the species. For sexually reproducing
organisms, a species consists of the various populations of individuals that are potentially capable of
breeding with each other and producing offspring that are not at such a disadvantage that they in turn
cannot breed successfully. The species itself is identified by two latinized names, the genus name
followed by the specific epithet, as for example in the scientific name for the Sacred Ibis, Threskiornis
aethiopicus.
Acceptability of names is governed by codified rules, especially the rules of priority—the first valid
name proposed for an organism is the name everyone is expected to use. Determining the right name to
use is sometimes but not always straightforward. Sometimes what constitutes the original description is
debatable; species repeatedly have been reassigned among genera as new information and opinions
about their relationships emerge. So most species have gone by various names since their discovery. In
each species account, the scientific name and information on the source of the original description are
given, as is the identity of the taxonomist credited with naming the bird. This attribution is placed in
parentheses if the genus to which the species is assigned has changed since description.
Unlike scientific names, common names vary from place to place and from language to language.
There has been some effort to standardize common names in various geographic regions, where
technical committees deliberate and provide lists of recommended usages. In each species account, we
make an effort to provide the reader with some of these names. There is, however, no 'right' or 'wrong'
common name, and some species have a score of names in several languages. The English names we
use, we recommend for their clarity within context of the stork-ibis-spoonbill group worldwide.
Over wide geographic areas populations of some species may diverge into recognizably different
kinds of organisms but yet be fully capable of interbreeding. These distinguishable geographic entities
are often called races or subspecies. Subspecies may be named, should it be useful to do so. For example
the Sacred Ibis of Africa is Threskiornis aethiopicus aethiopicus, whereas the isolated birds of Madagascar
are Threskiornis aethiopicus bernieri. In most cases, populations are not named, even though we can expect
variation to occur among populations scattered over wide areas of the globe.
What population should be considered a species and what a subspecies is often unclear, and so may
be contentious. Interbreeding potential is what marks a species. Members of two different species would
not interbreed if they occurred together; members of the same species or two subspecies of the same
6 Classification

species would; however, since they do not occur together,

proof of their compatibility is usually lacking. To be con-
sidered a subspecies, a population must be distinctive yet
potentially interfertile with other subspecies and capable of
producing offspring that are not at a disadvantage in nature.
(A species having more than one subspecies is called polyty-
pic.) One should keep in mind that the more fundamental
question of interest is not whether a population has a sub-
species name, but rather what are the biological impli-
cations of the variation of characteristics among popu-
lations.
Species that are similar, and obviously closely related,
that are found in disjunct ranges are called allospecies and
considered members of a superspecies. The Sacred Ibis,
Oriental White Ibis, and Australian White Ibis are mem-
bers of a superspecies, in our classification scheme.
The reproductive isolation of individuals of two species
results from differences in morphology, behaviour, biochem-
istry and, ultimately, genetics. These all provide criteria by
which we too can identify various species, and also the
criteria upon which are based the higher classifications,
which attempt to express the genetic similarities that derive
from sharing common ancestors.
The genus name is a low-level way to provide an indi-
cation of relationships among species. Species that share the Figure 5. An American White Ibis in breeding plumage,
same genus name are more closely related to each other with gular-sac inflated (southeastern USA).
than they are to species in other genera. That is, species
sharing a genus name also share a common ancestor, and a Figure 6. American White Ibis. Immature (left) and
genetic similarity, not shared with other species. adult (southeastern USA).
 Classification 7

Higher-level classification proceeds in the same manner, arate these species from other birds, some of which may or
with related genera being grouped into families, families may not be included in the order. This deficiency has led to
into orders, orders into classes, classes into phyla, and phyla considerable debate, over the past 50 years, as to which
into kingdoms. To show finer relationships, primary taxa birds should be included in the Ciconiiformes (see Sibley
can be further subdivided into higher and lower groups, and Ahlquist 1972 and 1990 for further discussion). Storrs
such as suborder (below order), subfamily and tribe (below Olson (1979) concluded that it was an artificial assemblage.
family) or superspecies (above the species). In some Alan Feduccia (1980) has suggested that resolution of the
detailed classification schemes, there may be many more phylogenies of the groups currently included in the Ciconii-
taxa used to show details of relationships. Above the species formes remains one of the major challenges of contemporary
level, classifications become increasingly a matter of pro- evolutionary ornithology. Either way, the use of biochemical
fessional opinion; although modern techniques of biochemi- characteristics has gone a long way to increasing our confi-
cal genetics and cluster analysis can make the process of dence in the close relationships among birds in the tradi-
seeking similarities more objective, questions of convergence tional ciconiiform groups.
nevertheless remain. The concept of a ciconiiform taxon has a long and dis-
tinguished history. T.H. Huxley (1867) suggested a group,
which he called Pelargomorphae, that included the ibises
STORKS, IBISES AND SPOONBILLS and spoonbills, storks, herons and flamingos. This was
Who, then, are the storks, ibises and spoonbills? According the basic view of Gadow and of Wetmore, who produced the
to currently accepted working classifications, storks, ibises classification scheme that for almost a century has been the
and spoonbills share a common classification down to the basis for the modern consensus of avian classification. Wet-
ordinal level. They are animals (kingdom Animalia), chor- more's (1960) Ciconiiformes was subdivided into four
dates (phylum Chordata), birds (class Aves), and wading suborders for the herons, the Shoebill, the storks, ibises and
birds (order Ciconiiformes). Which birds and which groups spoonbills, and the flamingos; these are the 'traditional
are additionally included in the order Ciconiiformes is, ciconiiforms'. The herons were further divided into two
however, a matter of considerable debate in the pages of the
scientific literature.
The storks, ibises, spoonbills, herons and others are
identifiable in part by their long legs and long necks, both
adaptations for foraging by wading about in shallow water.
These few characteristics of obvious ecological importance
may seem to be rather slim branches on which to hang a
taxonomic relationship. However, this intuitive grouping
has held up to repeated scrutiny since Linnaeus in 1758, and
we can be reasonably confident that the characteristics
represent a fairly robust adaptive strategy.
Other characteristics of the storks, ibises and spoonbills
relate to the adaptive strategy of being a 'wading bird'. Their
long necks have 15-20 vertebrae, and are capable of con-
siderable mobility, as would be expected to be able to feed.
Their tibiae and tarsi are unfeathered, and toes are
elongated, often slightly but never fully webbed, with the
lengthened back toe (hallux) also expected of a wading bird.
Some groups sport toenails specialized for feather mainten-
ance. They also have other characters in common (T.H.
Huxley 1867, Baird et al. 1884, Goues 1903, Witherby et al.
1939), such as similar sterna, palates and syringes, minute
intestinal caeca, elongated bills, long, broad and rounded
wings, and short tails. But other birds also have some of these
characteristics. They also share such common morphologi-
cal features as particular conformations of carotid arteries
and bony palate, which however are not unique to these
birds. The sexes are identical in plumage; but females are, to
a greater or lesser degree, smaller than males. Many species
nest in colonies and all have helpless (altricial) young that
adults feed in the nest.
In addition to the similarities, the groups of birds tra-
ditionally included in the order differ in aspects of behaviour
and morphology. It is, in fact, not possible to enumerate Figure 7. A Royal Spoonbill with crown feathers fully
obvious morphological characteristics that definitively sep- erect (southeastern Australia).
8 Classification
families, the typical herons (Ardeidae) and the Boatbilled
Heron (Cochleariidae). The storks (Ciconiidae) and the
ibises/spoonbills (Threskiornithidae) were in their own in-
dividual families, as was the distinctive Hamerkop (Scopi-
dae). Wetmore's classification, minus the flamingos, was
similar to that proposed about the same time by Verheyen
(1959) and was congruent with one of the first applications
of biochemical genetic data by Sibley (1960a,b).
Since then, other studies have been conducted, including
a proposed revision of the classification of the world's birds,
based on biochemical and other evidence, by Charles Sibley
and his associates (Sibley et al. 1988, Sibley and Ahlquist
1990, Sibley and Monroe 1990). These studies, in one sense,
produced a radically different view of the Ciconiiformes, a
more encompassing view that includes within the order
other groups. Many of these have previously been suggested
to be related in one way or another, but they were generally
retained in other orders. At the same time, taxa have been
removed from previously recognized orders, implying that
these were not natural assemblages. This expanded ciconii-
form order included, in addition to our groups, such groups
as the waders (i.e. shorebirds), gulls and terns, hawks,
grebes, tropicbirds, boobies, anhingas, cormorants, herons,
the Hamerkop, flamingos, pelicans, New World vultures,
frigatebirds, penguins, loons, petrels, and albatrosses.
Figure 8. A female Yellowbilled Spoonbill performing the
Greeting Display (southeastern Australia).
In another sense, these studies have been comforting in
that nestled within the expanded Ciconiiformes are the
herons, storks, ibises and spoonbills, which continue to be
perceived as being related at some level between family and
order.
It is useful to draw a distinction between a classification
and a species sequence (Bock 1990). A classification is a
three-dimensional tree of relationships that ideally reflects
the phylogeny (i.e. the evolutionary relationships) of the
group (Sibley and Ahlquist 1990). A sequence is an ordered
list of taxa, derived from a classification schema. A standard
sequence is a heuristic device that facilitates communi-
cation, and is not tampered with lightly (Mayr 1989, Bock
1990). In this monograph we have followed the standard
species sequence of Peters' (1931) Check-list (Kahl I979a,b,
J. Steinbacher 1979), to the fullest extent possible given the
new understandings of species limits adopted in this book.
Questions about Ciconiiform Taxonomy
All classification systems are really the expression of a series
of hypotheses that provide the basis for new questions and
new hypotheses (Sibley et al. 1988). Some of the principal
taxonomic questions exposed by proposed classifications
include: the relationships of the flamingos; the relationship
Figure 9. A Marabou Stork yawning at its nest (north-
western Kenya).
 Classification 9

of the herons to the other groups; the most appropriate

classification of the Hamerkop; the most appropriate classi-
fication of the Shoebill; the boundaries of the order with
respect to possibly related taxa such as pelicans and their
relatives, New World vultures, and hawks; and the family
and generic relationships among the various species within
the constituent groups of the traditional ciconiiforms.
The relationship of the flamingos has been an especially
controversial question, particularly because they show mor-
phological similarities to ciconiiforms, waterfowl and
waders. The question of interest in our context is whether
the similarities they show to other long-legged wading birds
might be due to convergence or to shared ancestry. Sibley
and Ahlquist (1972) concluded that flamingos were indeed
ciconiiforms and that flamingos linked the wading birds and
the waterfowl. Feduccia and Olson (Feduccia 1978, 1980,
Olson 1979, Olson and Feduccia 1980), on the other hand,
argue from palaeontological and other evidence that shore-
birds constitute the ancestral group of both the flamingos
and the waterfowl, and that none of these groups is particu-
larly closely related to the ciconiiforms. On the basis of their
studies of DNA, Sibley and Ahlquist (1990), however, con-
clude that flamingos are indeed relatively closely related to
other members of the heron-stork-ibis cluster of birds.
The herons are, by all accounts, closely related to each
other (Payne and Risley 1976). They have distinctive, Figure 10. Abdim's Storks at their nest on a rocky island
narrow feather tracts and patches of powder-down feathers, in Lake Shala (southern Ethiopia).
a completely feathered head, seasonal development of
elongated display plumes, a narrow skull, a long, straight the order to ibises and storks (Sheldon 1987). More specifi-
bill, a modified 6th cervical vertebra that allows the neck to cally, this evidence suggests that herons, the Hamerkop,
be held in an S-shape in flight, a comb-like (pectinate) flamingos, ibises, storks, the Shoebill and pelicans are de-
middle toenail, and young with a single down coat. Hancock scended from a common ancestor (Sibley and Ahlquist
and Kushlan (1984), in a companion volume to the present 1990).
monograph, provided a synthesis of the biology and classifi- The Hamerkop (Scopus umbrettd) is a small bird, lacking
cation of the group, the latter mostly derived from the the long legs and neck characteristic of the other traditional
studies of Payne and Risley (1976) and Payne (1979), ciconiiforms. We have found that it flies, perches, feeds,
although retaining several traditionally generic allocations nests, and engages in courtship, all in distinctive ways
pending further study. (Kahl 1967b, 1979c). As noted above, biochemical evidence
One of the longstanding issues regarding the heron group (Sibley et al. 1988) suggests that it is related to the herons,
relates to the distinctive Boatbilled Heron (Cochlearius coch- storks and ibises, but it certainly is not a true stork. Because
learius). Hancock and Kushlan (1984) suggested that it of its distinctiveness, we have chosen not to treat the
should be grouped with the night herons. Sheldon (1987) Hamerkop in the present monograph.
more recently provided biochemical evidence for a subfami- The Shoebill (Balaeniceps rex) shares anatomical charac-
lial relationship between the Boatbilled Heron and tiger- teristics with the true storks, including a distinctive tubular
herons, and no distinction between day and night herons. ear bone (Feduccia 1977) and a Bill-clattering Display
Despite these finding, the linear arrangement of taxa by (Kahl 1972e). It also differs in other respects. For example,
Payne and Risley (1976) and the overall integrity of the we found that it did not excrete on its legs for cooling (Kahl
heron family continues to remain intact. If anything, recent 1967a) and its chromosomes differ from those of the true
studies have tended to find the Boatbilled Heron to be less storks (De Boer and Van Brink 1982). It shows many
distinctive from the other herons than previously thought. similarities to pelicans (Cottam 1957, Feduccia 1977, Saiff
However, the relationships of herons to other wading 1978, Olson 1979, 1985b). Sibiey and Ahlquist (1972) con-
birds has not been so well settled. Olson (1979) suggested sidered the species to be a ciconiiform, and Feduccia (1977)
that they show few morphological similarities to the other considered it to be a stork. Kahl (1979a) and Olson (1985b)
traditional ciconiiforms and are the only currently success- assigned it to its own family adjacent to the true storks.
ful representatives of an early radiation of crane-like birds Sibley and Ahlquist (1990) proposed that, within their
not clearly related to modern storks or ibises. He considers much expanded Giconiiformes, it is most closely related to
the family to be one of uncertain affinity (Olson 1985a,b). pelicans, and with the pelicans is placed in a superfamily
Biochemical genetic evidence continues to support their between the superfamily that includes the ibises and the one
ciconiiform heritage and a relationship at some level within that includes the storks and New World vultures. Because of
10 Classification
Figure 11. A Lesser Adjutant Stork feeding its young
(northeastern India).
its historical identification with storks, we have chosen to
include it in this monograph.
Although the boundaries of the order Ciconiiformes con-
tinue to be debated, there emerges a clear consensus regard-
ing the storks, ibises and spoonbills. From the morphology-
based classifications of Huxley and Wetmore, through the
various behavioural studies such as some of our own, to the
biochemical studies of Sibley and others, these three groups
of birds are repeatedly found to be related. Analysis of DNA
hybridization similarities indicate that herons, the Hamer-
kop, flamingos, ibises and spoonbills, storks, New World
vultures, pelicans, and the Shoebill are more closely related
among themselves than they are to other groups of birds and
so are descended from a common ancestor.
Sibley and Ahlquist (1990) describe well the probable
history of this group. Each is a result of a radiation exploit-
ing an unoccupied niche, each showing its own way of life.
Each flourished, and spread widely, retaining character-
istics of their common ancestry in their morphological and
genetic similarities.
In this monograph, continuing our evaluation of the bi-
ology of these groups that began with The Herons Handbook
(Hancock and Kushlan 1984), we summarize the biology of
the storks, the ibises, the spoonbills and the Shoebill.
Relationships within the Storks
The typical storks are a distinctive group sharing a number
of characteristics, including long legs with half of the tibia
bare, relatively short toes with small webs, 12 tail feathers,
12 primary feathers (the 12th being minute and/or lacking
in the wood storks and openbills), bare portions on the
head, feathered oil gland, a common musculature of the
pectoral associated with soaring flight (Vanden Berge
1970), a stout bill, predominantly simple black and white
plumage colours, young with two down coats, large air sacs
under the neck skin, and a lack of powder down. They also
have behaviours in common, including cooling their legs by
defecating on them (Kahl 1963b), a trait they share with the
New World vultures (Ligon 1967). Olson (1985b) and
Sibley et al. (1988), and others before them, suggested that
they are closely related to the New World vultures, and
Sibley and Ahlquist (1990) included the vultures in the
same family as the storks, remarking that they had been
taxonomically misplaced for two centuries.
The most recent classifications of stork species are based
on behaviour (Kahl 1979b), anatomy (D.S. Wood 1983),
and a combination of the two (D.S. Wood 1984). These
studies have grouped the species in fewer genera than was
previously the case (Kahl 1972e). We draw from the results
of both sets of studies in recognizing two higher level div-
isions of the true storks, the wood storks and the openbills in
the tribe Mycteriini, and the typical storks, giant storks and
adjutants in the tribe Ciconiini. We have chosen to use
tribes in subclassifying the storks, rather than the more
traditional rank of subfamily, because the proper taxonomic
rank of the typical stork group remains under discussion as
does the appropriate positioning of the Shoebill and New
World vultures. Sibley et al. (1988) classify the traditional
storks as a subfamily, in which case tribe would be the next
lower taxon. In the present monograph, we continue the
more traditional approach of considering the storks to rep-
resent the family Ciconiidae.
The stork tribe Mycteriini includes four wood storks
(Mycteria) and two openbill storks (Anastomus). All locate
food primarily by touch rather than by vision, and have bills
especially, and even uniquely, suited for this behaviour.
The wood storks have large, cylindrical, but slightly
down-curved bills, which they place in the water and hold
open for fish to swim into. They also use special movements,
such as pumping their feet or opening and closing their
wing, to increase the likelihood of a fish entering the bill.
Mycteria have distinctive displays including aerial Flying

Around, a Gaping Display, and Display Preening (see sub-
sequent chapters for descriptions of these and other behav-
iours). Courtship includes a gaping and snapping of the bill
but not the Bill-clattering characteristic of other storks. The
differences among the Mycteria storks are minor, consisting
primarily of plumage and soft part coloration (Kahl 1972b).
It is interesting to note that one of the wood storks, the
Yellowbilled Stork, is called Mycteria ibis. Through the
vagaries of nomenclatural history, the rules of priority result
in the name 'ibis' being given not to an ibis, but to a stork
(Mathews 1913).
The openbills have bicurved bills that leave a central gap.
Where they come together at the tip, the curved mandibles
form a forceps with which the stork pincers its preferred and
almost exclusive prey, freshwater snails. The two species are
markedly different in colour, Anastomus lamelligerus being
black and Anastomus oscitans being mainly white. The two
possess an Advertising Sway behaviour that seems to be
unique (Kahl 1972d); during copulation the male shakes his
head rather than clattering his bill.
In the tribe Ciconiini, we include 13 species in the genera
Ciconia (typical storks), Ephippiorhynchus and Jabiru (giant
storks), and Leptoptilos (adjutants). D.S. Wood (1984), argu-
ing on the basis of his quantitative analysis of character
distance, deviated from Kahl's (1972e) classification by
Figure 12. Asian Openbill Storks sunning (northern India).
Classification 11
including the genera Ephippiorhynchus and Jabiru in a tribe
with the typical storks. This reflects the undoubted affinities
between the genera Ciconia and Ephippiorhynchus. However,
the Jabiru also shares characteristics with the adjutants, and
is somewhat of an intermediate between the other two giant
storks and the adjutants. In that the classification scheme
should reflect that intermediacy, it seems to us that a more
natural classification is achieved by grouping all four genera
into one tribe, Ciconiini.
The typical storks, members of the genus Ciconia^ have
straight or slightly recurved bills; the head and neck are
feathered, but the skin around the eye (lores) is bare. A
display, the Head-shaking Crouch, performed by the male is
found uniquely in the genus Ciconia (Kahl 1971f, 1972c).
Bill-clattering and whistling vocalizations are characteristic,
especially in courtship. We have concluded that the Black
Stork is the most primitive and the White Stork the most
advanced in the genus Ciconia (Kahl 1972c). Abdim's seems
closely related to the Black Stork, both behaviourally and
morphologically. The Woollvnecked Stork of Africa and
Asia and the Maguari of South America share a number of
characteristics, including a forked black tail. We here recog-
nize the Storm's Stork (Ciconia stormi) as a separate species
from the Woollynecked Stork and the Oriental White Stork
(Ciconia boyciana) as a separate species from the White Stork.
12 Classification

Fossils of this group are known, including both extant and following WJ. Bock (pers. comm. 1990) in his advocacy of
extinct species of Ciconia as early as the middle Miocene and retaining the well-used name, as well as by priority, we
as late as the late Pleistocene in Africa (CJ.O. Harrison continue to use 'Threskiornithidae'.
1980). The notable differences in bill morphology have long
In the giant stork genera Ephippiorhynchus and Jabiru, a resulted in the recognition of two subfamilies, the Threskior-
Flap-dash Display is used by pairs on the feeding grounds. nithinae including the ibises, having a narrow downcurved
The Blacknecked and Saddlebill Storks are similar in many bill with a flattened tip, and the Plataleinae including the
ways to the typical storks. The Jabiru shares an inflatable spoonbills with a flattened bill. The two bill types correlate
throat sac with the adjutant storks. Additionally, the Sad- with the predominant non-visual feeding modes adopted by
dlebill Stork has a vascularized chest patch. Because of its the two groups, Probing and Head Sweeping, respectively
intermediacy between the typical storks and the adjutants, (Kushlan 1978a). The various species have a number of
unlike D.S. Wood (1984), we have retained it in its own characteristics in common, especially a distinctive slit-like
genus. The adjutants (Leptoptilos) all feed to some degree as cranial morphology (called schizorhinal), which they share
scavengers, have massive bills, and lack normal contour- with waders (perhaps by convergence) and which is an
feathering on the head and neck, both adaptations for eating adaptation to allow upper bill movement. They also have a
large carrion. They are unique in giving a loud squealing or distinctive musculature and sternum, 11 primary feathers
mooing vocalizations during courtship (Kahl 1966a, 1970, ( l l t h minute), 12 tail feathers, long legs with lower half of
1972a). The large inflatable throat sac is an obvious charac- tibia bare, loss of feathers on the head and even neck,
teristic of these storks. distinctive ornamental plumes, and changeable soft-part
colours. They lack powder-down patches, have a cupped
middle toenail undoubtedly used for feather maintenance,
Relationships within the Ibises and Spoonbills and a grooved mandible that functions in removing se-
Ibises and spoonbills are medium-sized birds that have been cretions of the supraorbital salt gland. They have down on
recognized as being related since at least the mid 1800s both feather tracks and non-feathered portions of the skin
(Nitzsch 1840). They are almost always grouped at the (apteria). An interesting aspect of the biology of ibises is the
family level (Sibley and Ahlquist 1972, 1990). The most variability of chromosome structure among some species, a
widely used name for the group is the Threskiornithidae very unusual trait among birds (including storks) in which
(Eisenmann et al. 1984). It has been argued that by priority chromosomal uniformity is the rule (Capanna et al. 1982).
the more appropriate name is Plataleidae (K.E. Campbell In addition to the traits they share with storks, the ibises
1986, A.R. Phillips 1986, Olson et al. 1986). However, and spoonbills share characteristics with the waders, tra-
ditionally classified in the order Charadriiformes. Olson
(1985b) included ibises and waders in the same order.
However, a relatively close relationship is not supported by
Figure 13. A Woollynecked Stork shading its young at DNA evidence (Sibley and Ahlquist 1990).
the nest (northern India). The ibises and spoonbills have been relatively little stud-
ied in the field because they are hard to observe and the
various species are well scattered around the world. Ex-
tremely little is known about certain rare or inaccessible
species.
The ibis subfamily Threskiornithinae consists of 23
species distributed among 14 genera. It is likely that the
present classification does not adequately represent re-
lationships among the species. However, taxonomically use-
ful anatomical trends among the species are not readily
apparent. It is also an old group, dating back to the Mio-
cene, about 25 million years ago. The earliest fossils are
assignable to a still living genus, the widespread Plegadis
(Plegadis peganus and Plegadis pharangitus—Olson 1981).
Species representing two other living genera are known from
the Pliocene of South Africa, Geronticus apelex and Threskiornis
aethiopicus, the bones of the latter being indistinguishable
from the modern form that occurs in the area today (Olson
1985a). Extinct species of present genera (Theristicus wetmorei
and Eudocimus peruvianus) are known from the Pleistocene in
South America (K.E. Campbell 1979). Especially intriguing
are subfossils of flightless ibises, Apteribis glenos, from the
islands of Maui and Molokai in the Hawaiian Islands and
Jamaica (Xenicibis xympithecus) (Olson and Wetmore 1976,
Olson and Steadman 1977, 1979, Olson and James 1991),
 Classification 13

which indicate the occurrence of very recent extinctions of

particularly vulnerable species of ibis. The Hawaiian forms
were undoubtedly lost through the hand of man.
Relationships among the ibis genera are not well under-
stood. The glossy ibises (Plegadis], represented so anciently
in the fossil record, today have a nearly worldwide distri-
bution, the only ibis genus to do so. The Glossy Ibis itself
(Plegadis falcinellus] is found on all inhabitable continents
and may be a relatively recent invader of the New World.
The Whitefaced Ibis is the South American and western
North American form, whereas the Puna Ibis is a South
American montane form.
From available fossil and osteological evidence, it appears
that the genus Eudocimus is closely related to the glossy ibises
(Mayr and Short 1970, Olson 1981). The genus has tra-
ditionally been considered to include two forms, the Ameri-
can White Ibis (albus) ranging from North America to
northern South America and the Scarlet Ibis (ruber) found in
northern South America. Because their recent studies
demonstrated considerable successful interbreeding of red
and white birds in a natural zone of overlap, Ramo and
Busto (1987) concluded that, by definition, the two forms
must be considered the same species. This conclusion is
supported by other evidence, not the least being that the
presence of the carotenoid pigment canthaxanthin, which
accounts for the colour of the red form, is unreliable as a
taxonomic characteristic (Sibley and Ahlquist 1990).
South America is a centre of ibis radiation. Most species
there are at this time assigned to unique genera, reflecting
the dearth of information available on extant and fossil
South American forms, lack of taxonomic study of the sub-
family as a whole, and the probable age of the group.
Theristicus is here considered to include a single polytypic
species. The genera Phimosus, Cercibis, Harpiprion, and Mesem-
brinibis are also represented by single species. Sibley and
Ahlquist (1990:852) present the intriguing result of one
experiment: that there may be a split within the family
between the South American genera (Mesembrinibis, Theristi-
cus, Cercibis, Eudocimus} and the Old World ibis genera and
spoonbills. This deserves further study.
The sacred ibises (Threskiornis] are widespread in the Old
World. Once divided into four species, they were combined
into two species and many subspecies by Holyoak (1970).
More recent studies of their behaviour, structure, and
chromosomes suggest that three species exist (Cramp 1977,
De Boer and Van Brink 1982, Mees 1982, Lowe and
Richards 1990). In this monograph we restore the division
of this genus into four species, three of which are allospecies.
De Boer and Van Brink (1982) suggested that the Austra-
lian White Ibis (Threskiornis molucca) may be the ancestral
form from which the other two species were derived.
Several species of ibis of Africa, Madagascar and the

Figure 14. A Strawnecked Ibis in breeding plumage

(southeastern Australia).

Figure 15. Black Storks at their nest; male nearest the

camera (southwestern Poland).
14 Classification
Figure 16. A Shoebill in a grassy marsh adjoining Lake
Kyoga (north-central Uganda).
Middle East are included in the genera Bostrychia, Lophotibis,
and Geronticus. It is likely that these are fairly closely related
to each other (Fry et al. 1985). Three genera of ibises are
found only in Asia, Pseudibis, Thaumatibis, and Nipponia.
Olson (1985b) suggested that Pseudibis is anatomically very
similar to and possibly congeneric with Geronticus. Although
the Giant Ibis has most recently been considered to be a
Pseudibis (Holyoak 1970), Olson (1985b) has cautioned
against removing this distinctive, large ibis from its special-
ized genus without anatomical study. We have followed this
sensible advice and restore it to its separate genus.
The spoonbill subfamily Plataleinae has a cosmopolitan,
although somewhat patchy, distribution. Four of the Old
World species are closely related, and largely replace each
other geographically. We here identify the Royal Spoonbill
(Platalea regia} as a separate species, based on our recent
studies of its behaviour and morphology, which differ from
the Eurasian species (Kahl 1988a). The specific distinctive-
ness between the Royal and Blackfaced spoonbills (Platalea
minor) also needs scrutiny. The New World Roseate Spoon-
bill (Platalea ajaja) and the Australian Yellowbilled Spoon-
bill (Platalea flavipes) are more distantly related to the other
spoonbills, and may be most closely related to each other
(Kahl 1988a).
Species Sequence of the Storks, Ibises
and Spoonbills
Family CICONIIDAE—-the true storks
Tribe MYCTERIINI—the wood storks and openbills
Mycteria americana — American Wood Stork
Mycteria cinerea — Milky Stork
Mycteria ibis—Yellowbilled Stork
Mycteria leucocephala—-Painted Stork
Anastomus oscitans—Asian Openbill Stork
Anastomus lamelligerus — African Openbill Stork
Tribe CICONIINI—the typical storks, giant storks and adjutants
Ciconia nigra — Black Stork
Ciconia abdimii—Abdim's Stork
Ciconia episcopus—Woolly necked Stork
Ciconia stormi—Storm's Stork
Ciconia maguari — Maguari Stork
Ciconia ciconia — White Stork
Ciconia boyciana—Oriental White Stork
Ephippiorhynchus asiaticus — Blacknecked Stork
Ephippiorhynchus senegalensis—Saddlebiil Stork
Jabiru mycteria— Jabiru Stork
Leptoptilos javanicus — Ltsstr Adjutant Stork
Leptoptilos dubius —Greater Adjutant Stork
Leptoptilos crumeniferus — Marabou Stork
Family BALAENICIPITIDAE — the Shoebill
Balaeniceps rex — Shoebill
Family THRESKIORNITHIDAE — t h e ibises and spoonbills
Subfamily THRESKIORNITHINAE —the ibises
Eudocimus ruber—American White Ibis
Phimosus infuscatus—Barefaced Ibis
Plegadisfalcinellus—Glossy I bis
Plegadis chihi—Whitefaced Ibis
Plegadis ridgwqyi — Pun& Ibis
Cercibis oxycerca — Sharptai/ed f bis
Harpiprion caerutescens—Plumbeous Ibis
Theristicus caudatus—BufFnecked Ibis
Mesembrinibis cayennensis—Green Ibis
Bostrychia kagedash — Hadada Ibis
Bostrychia carunculata — Wattled Ibis
Bostrychia olivacea—OVive Ibis
Bostrychia rara — Spotbreasted Ibis
Lophotibis cristata — Madagascar Crested Ibis
Threskiornis aethiopicus—Sacred Ibis
Threskiornis melanocephalus — Oriental White Ibis
Threskiornis molucca—Australian White Ibis
Threskiornis spinicollis — Strawnecked Ibis
Geronticus eremita — Waldrapp Ibis
Geronticus calvus — Bald Ibis
Pseudibis papillosa—Black ibis
Thaumatibis gigantea — Giant Ibis
Nipponia nippon—Oriental Crested Ibis
Subfamily PLATALEINAE—the spoonbills
Platalea leucorodia — Eurasian SpoonbiJJ
Platalea regia — Royal Spoonbill
Platalea minor—Blackfaced Spoonbill
Platalea alba — African Spoonbill
Plataleaflavipes~ Yellowbilled Spoonbill
Platalea ajaja — Roseate Spoonbill
 Conservation

H
ow unfortunate it is that large water birds are particularly subject, either intentionally
or otherwise, to continual attack from the human race. They are hunted, they are eaten,
they are harassed, their feeding habitat is altered, their nesting sites are destroyed. On
the other hand, they are also appreciated, for their size, their beauty or ugliness, or for
the place many hold in the culture and history of a people.
In our studies of the herons of the world, we were constantly amazed at the resilience of members of
that family (Hancock and Kushlan 1984). In our work on the present volume, we found that for storks,
ibises and spoonbills the story is a very different one. These birds have come under much the same
pressures as have herons, but for reasons not quite clear, except as explicated for an individual species,
they have responded poorly to the changes imposed upon them. A result is that many species of this
diverse group of birds are at, or near extinction, over much of their former range (W.B. King 1981,
Collar and Andrew 1988).
In many areas, these bird populations are suffering from a combination of pressures including
deliberate killing, deforestation, wetland drainage, use of intensive modern farming methods, and the
accumulative effects of pesticides (J.F. Parnell et al. 1988). These problems seem universal, though the
degree to which they apply varies according to the awareness and concern of the human inhabitants of
the region. In fact, almost invariably for these large birds, it is the attitude of the local human
population that decides whether a species survives in its midst.
Solutions are by no means unclear, although they may be difficult to put into practice; J.F. Parnell et
al. (1988) discuss these. Habitat improvement is required to reverse the impact of wetland destruction
and alteration. In many cases improvement will require direct manipulative management of water,
vegetation and breeding sites in compensation for loss of natural feeding and nesting sites, as well as
monitoring and protection of existing functioning ecosystems (Kushlan 1979b, 1983, 1987, 1989a,b).
More general environmental improvements are also required, especially regarding the long-term effects
of chemical contaminants. It has been noted that 'People control is a foremost aspect of colonial
waterbird management' (J.F. Parnell et al. 1988), and control of disturbance, including both passive
disturbance and hunting, is needed, especially at colony sites. In some cases it is necessary to consider
management to reduce natural interactions, such as predation and competition, for crucially endan-
gered species. Multispecies fishery management is required in those areas where the needs of these birds
are at odds with those of human fisheries. In some cases, the enhancement or re-establishment of a
population may require reintroduction of birds, from wild or zoo stocks, and the artificial establishment
of suitable nest or colony sites (e.g. Hirsch 1978a,b). Also critical are additional research on limiting
factors and increased emphasis on education, especially of rural peoples.
To be most effective, these management and conservation activities should be enacted on a regional
basis. Luthin (1987), in discussing the conservation needs of storks, noted that a species' conservation
concerns will often differ within its overall range. Also, because more than a single wading birds species
may be dependent on regional resources, it is often within this multispecies context that management
plans should be drafted. These plans, single-species or regional, are essential to define and guide
conservation activity.
In some parts of the world, mankind is belatedly attempting to halt the destruction of forest and
wetland habitat in which the majority of storks, ibises and spoonbills live. But it is not clear if it is too
late for some populations.
Endangered means that a species or subspecies is in danger of extinction, and its survival is unlikely if
the causal factors continue to operate (Collar and Stuart 1985). Populations that are decreasing may be
16 Conservation
Figure 17. A Milky Stork flying over the nesting colony (northwestern Java, Indonesia).
considered vulnerable if further declines could make them
endangered. Some species are vulnerable because of their
rarity, which may or may not be due to human intervention.
We consider five species and two subspecies to be endan-
gered, and six species to be vulnerable. These species en-
compass over 20% of the world's species of storks, ibises and
spoonbills.
Amongst the storks, a number of species are endangered
and many more are decreasing in numbers in some or all of
the areas they inhabit (Luthin 1987, Collar and Andrew
1988). The Storm's Stork is a vulnerable species that has
never been common and is seldom seen. Until recently we
were not even sure of its status and so may never fully
understand its biology before it becomes extinct. Other
species are rare because of man, the Lesser Adjutant (vul-
nerable) and Greater Adjutant (endangered) Storks have all
but disappeared over most of their range and are now in
grave danger. The vulnerable Milky Stork remains secure
only in the Indonesian archipelago. The Oriental White
Stork and Black Stork, both vulnerable species, are increas-
ingly confined to isolated breeding sites.
Ibises and spoonbills have a number of species at risk
(Manry 1986). Of the six species of spoonbill, the Black-
faced Spoonbill is the most endangered, having retreated as
a breeding bird to a few islands off the North Korean coast
or perhaps to other unknown sites yet to be discovered. The
endangered Giant Ibis may just be hanging on in isolated
areas in Cambodia and perhaps Vietnam. The endangered
Davison's race of Black Ibis now occurs only in Vietnam
and possibly Indonesia. The endangered Waldrapp Ibis has
now gone from Turkey, appears to be rapidly decreasing in
Morocco, and seems to be on the verge of becoming extinct
(Hirsch pers. comm.). The highly endangered Oriental
Crested Ibis is now known to occur in the wild only in
Qinling Shan in south Shaanxi, China, where the known
population is in the low dozens. The continued existence of
the dwarf Sao Tome Island race of the Olive Ibis has only
recently been confirmed, but the situation is extremely pre-
carious for this endangered race. The once widespread
Madagascar Crested Ibis is vulnerable due to habitat
changes that have been occurring in Madagascar.
Everywhere pressures of human population have affected
even the widespread species, and as suitable environments
shrink so the pressures on all species increase. Regional
decreases in numbers are documented in most species for
which such data are available. It is likely that in the last 50
years that populations of most storks, ibises and spoonbills
have been reduced by at least a half, and in many species by
more. The extent of these reductions vary regionally.
Bird populations in North America are probably the best

known in the world. Trend in numbers and distribution
suggest that considerable changes in ranges and populations
have been underway for many decades. Perhaps the greatest
decreases are in the Florida Everglades, where once thriving
populations of American Wood Storks and American White
Ibises have been decreasing to the point of local extinction
(Frohring et al. 1988). The desperate position of the Ciconii-
formes has developed with great rapidity. On the other
hand, these species still maintain sizable populations else-
where in North America and into South America. Within
the United States the conservation and management needs
are becoming well recognized (J.F. Parnell et al. 1988), but
many countries have not the resources available in the USA
to fight for the survival of decreasing species.
Over much of South America, populations of all species
remain high (or unknown). Even where historical decreases
have occurred, there is evidence that populations may be
returning. However in South America, apart from a few
individuals and isolated concerned government or private
bodies, conservation of habitat or species is not yet under-
way.
In Europe the protection of species by concerned govern-
ment and private bodies is holding back the deterioration,
but species such as the White and Black Storks are falling in
numbers annually. Eurasian Spoonbills are confined to a
Figure 18. A Bald Ibis (northeastern South Africa).
Conservation 17
few sites in western Europe. The former indifference, or
active persecution, in western Europe has, except in Medi-
terranean regions, belatedly been replaced by a growing
concern, manifesting itself into sometimes ineffective but
now increasingly professional attempts to protect such
species on their feeding and breeding grounds. Quite re-
cently, however, it has become clear that in eastern Europe,
pollution created by ill-conceived and inefficient industrial-
ization is on a much larger scale than was previously rea-
lized. Some areas, it is thought, may never recover from the
environmental damage, but all require attention.
In Africa, the picture is probably the most favourable as
most species remain widespread and even common (Collar
and Stuart 1985). North of the Sahara, the Waldrapp Ibis is
fast on its way to extinction. It is well to note that such local
extermination is not new in the area. The ancient Egyptians
mummified Sacred Ibises by the hundreds of thousands,
perhaps over a million, undoubtedly reducing the popu-
lations that probably were finally extirpated by land use
changes (Goodman 1988). Modern times have produced oil
spills and the catastrophic effects of recent wars have yet to
be evaluated. Historically the peoples of subSaharan Africa
have shown tolerance to creatures around them, but in-
creases in human population have begun to change the face
of this continent, and the need to feed increasing numbers
Figure 19. A female Blacknecked Stork foraging (north-
ern Australia).
18 Conservation

of people has, together with ill-conceived development

schemes often funded by western agencies, put pressures on
all species of storks, ibises and spoonbills (Collar and Stuart
1985). Island forms in the Atlantic and Indian Oceans
continue to be at great risk.
In Asia the picture is very bad (Bain and Humphrey
1980, Hancock 1989). Here many wading bird species are
very near extinction over much of their former range. Only
in parts of India and in the, as yet, undeveloped areas of the
Indonesian archipelago do some of the former abundant
species survive in any numbers. The reasons for such a
dramatic change, which has occurred since the end of the
Second World War —a matter of about 45 years — are not
hard to find. The elimination of endemic diseases and the
resultant population explosion combined with a huge in-
crease in the standard of living of many, though by no
means all, its peoples have meant a rapid and urgent need to
exploit all its natural resources (D.A. Scott and C.M. Poole
1989). Southern and eastern Asia comprises only 14% of the
world's land surface, but support at the present time 56% of
the world's human population. Furthermore this huge
population is expanding at about the rate of 55 million per
annum (D.A. Scott and C.M. Poole 1989).
Also in Asia, not only has continuous warfare in many
areas destroyed much of the formerly unexploited natural
habitat, but the demand for food under such circumstances
has meant the killing of large birds for food. Weapons are
now readily available for shooting, and in many areas no
feeding bird is safe from the gun, and no breeding bird safe
from the trap. There has been a breakdown in the tradi-
tional religious beliefs in some nations, notably China, such
that temple grounds and monastries, which traditionally
provided protection for storks and ibis colonies, now seldom
do so today. An exception which still enjoys protection
in the temple confines is the Asian Openbilled Stork of
Thailand.
Today's awareness of the need for conservation measures
has barely touched some nations or regions of Asia. The
peoples of Pakistan, have a tradition of hunting which, using
modern weapons, has served to nearly exterminate all the
species of wading birds that bred in their land. War and
poverty in the Philippines have brought with them heavy
deterioration in the numbers of large birds. In Japan the
recent stirring of conscience there has come too late in the
day for most species.
The subcontinent of India provides, perhaps, the clearest
example of how religious and traditional tolerance has
meant the survival of storks, ibises and spoonbills, and other
resident and migratory species of large water birds. In areas
where such ancient customs either do not pertain or have
been lost beneath the struggle to survive, such species have

Figure 20. Jabiru Storks at the beginning of a copulation,

with neck-bands a fiery red (northeastern Argentina).

Figure 21. A female Saddlebill Stork at its nest with three

nestlings about 45-50 days old (western Kenya).
 Conservation 19

been exterminated. An example is the Black Ibis, which has Popular programmes, environmental education, and in
its stronghold in Gujarat, a state composed mainly of toler- country and out of country workshops for wildlife pro-
ant, strict Hindu peasants and landowners. Here this fessionals may be used with success. Without such edu-
species is quite tame, feeding around village ponds and cation, the best laid conservation scheme will come to
nesting near to houses and other occupied buildings. But in naught.
states where hunting and trapping are still carried out, it is
only in the nature reserves that the Black Ibis survives.
Even so, in Gujarat, such tolerance has not saved the Black-
necked Stork. Drought conditions have dried out wetlands,
and increasing demand for firewood has resulted in the
destruction of trees on a vast scale, so that this bird has lost
its traditional nesting sites, with the result that no breeding
pairs remain there.
Australia is the area of the world that retains its several
species in greater abundance than perhaps any other. The
Blacknecked Stork is at the southernmost limit of its range
in Australia and New Guinea, having been almost totally
wiped out in the rest of Asia apart from a few pockets of
birds in India and perhaps Cambodia and Vietnam (Han-
cock 1989).
Conservationists and conservation organizations in India,
Africa and Asia are struggling with the problems they face.
Private organizations such as Interwader in Asia are survey-
ing the shrinking wetlands and recommending methods of
protecting them. The ICBP is donating funds for work in
many parts of the world. Brehm Fond has sponsored sur-
veys and workshops on colonial bird conservation in Asia.
In spite of all the measures being taken, there is an
inevitability about the demise of some species. Given the
need to preserve genetic stocks represented by these species
and the hope that habitat restoration will allow reintroduc-
tion, it would seem likely that captive breeding remains the
only hope in some cases. The Waldrapp Ibis thrives in
captivity with some 500 birds, whereas in the wild the total
population may well be below 100. The Oriental Crested
Ibis numbers in China may be increased if a way can be
found to breed them successfully in captivity. Many storks
prosper under captive conditions and could be released in
the wild if suitable protected areas can be found for them.
The zoological parks of the world are attempting to breed
these species, and such efforts should be encouraged
( L u t h i n e t a l . 1986, R.E.Johnson et al. 1987a,b). Of course,
all proposed reintroductions require detailed consideration
and study (Kushlan 1980, Seward 1987).
The need for education is paramount to the success of any
conservation measures. The recent discovery of the few
colonies of Greater Adjutant Stork breeding in villages in
Assam heightens the need in that area (Saikia and Bhatta-
charjee 1990) As a scavenger the species requires a food
supply provided by man and his domestic animals, and does
not appear to thrive in nature reserves where man is
excluded. Conservationists, scientists, farmers and local
landowners do not mix happily and readily together, and
great care will be required if useful and productive under-
standing is to be reached in Assam, and indeed in many
other areas where man, stork, ibis and spoonbill live in close
proximity.
Similarly, in South America, education efforts must be Figure 22. White Storks copulating on a rooftop nest
aimed at both the populace and government agencies. (southwestern Poland).
 Courtship and
Reproduction

I
n order to reproduce, storks, ibises and spoonbills must form pair-bonds, mate, build nests, lay,
incubate and defend their eggs, and rear their young at least until the chicks are able to fly
safely from the nest. Reproduction is a complicated affair, requiring in all these species the
cooperation of both parents.
A few species, such as the Black Stork, and the Hadada, Spotbreasted, and Black Ibises, nest
alone in pairs. These usually are found in tall trees, especially in forests. Most species nest in colonies.
Colonial species may nest on the ground, in reeds or in trees, depending on the species and the
environment. Perhaps the most unusual are those that nest on cliffs, including the Waldrapp, Bald,
Wattled and Buffnecked Ibises.
Timing of nesting differs among species and locations. Both photoperiod and environmental con-
ditions affecting food availability seem crucial in determining nesting timing. In the high latitudes,
nesting is decidely seasonal and is associated with migrations out of the nesting area in the local winter.
In the subtropics, nesting is often in the drying season. In the tropics, some species may nest nearly all
year round. In all cases nesting occurs during the local period of high food availability. In times of low
food availability, especially due to inappropriate rainfall or surface water conditions, nesting may be
postponed or may be skipped in a given year. The Waldrapp and American White Ibises are examples
(Smith 1970, Kushlan 1978b). The tight relationships of rainfall to nesting is particularly demonstrated
in Australia, the Florida Everglades and Africa (Carrick 1962, Urban 1974b, Kushlan 1989a,b).
Meeting and overcoming environmental and social constraints on reproduction are achieved through
the use of appropriate behaviours, some of which have been ritualized into displays that allow one bird
to communicate with another.

THE PAIR-BOND
The pair-bond is the key to successful nesting. Both members of the pair participate in building and
protecting the nest, incubating the eggs, and feeding and caring for the young until they fledge from the
nest site under their own power. It seems that in these species a single parent is insufficient to supply
simultaneously the protection and food needed to raise young.
Despite the importance to both individuals of maintaining a strong pair-bond to raise their young, it
is becoming increasingly clear that promiscuous mating behaviour, called extrapair copulation, is
extremely common in some species. Colonial nesting presents opportunities for promiscuity that are less
available when, as in most species, birds nest in widely separated nesting sites within defended
territories. The evolutionary advantage is clear: the promiscuous male increases its chance to sire
additional offspring while the female is receiving a bold and clearly successful male who might father
some of her young. But risks are also present, not the least of which is the potential loss of one's own nest
or eggs if they are left unattended, as we have seen in the American White Ibis (Kushlan 1973b).
An important question is whether extrapair copulations result in insemination, and this is a difficult
question to study in these birds. Another question is what are the strategies the male uses to defend its
mate against such intrusions, while also attending to its other duties such as supplying nest building
materials? In the Eurasian Spoonbill, most extrapair copulations occur while the resident male is away
gathering nesting material; and, perhaps as a response, males usually copulate with their mate just
 Courtship and Reproduction 21

before leaving them to go on stick-collecting trips (Aguilera according to their component body movements, without
1989, Aguilera and Alvarez 1989). Despite promiscuity, assigning or assuming motivation. For purposes of enhanc-
even if an intruder is caught in the act, the initial pair-bond ing clarity, increasing descriptiveness and reducing motiva-
usually persists, probably because the pair has little choice if tional inferences, we have renamed some of the terms cur-
they are to raise their young. rently found in the literature. These changes are noted in
Pair-bonds last for at least a nesting episode. If nesting is the accompanying table.
interrupted at an early stage, many species can nest again So that the named ritualized behaviours will stand out
during the same season. Individuals of some species may re- and be easier to locate in the text, they have been capital-
pair in successive years, or even remain paired year-round. ized. In general, using the same name for a behaviour in
The White Stork is famous for maintaining pair-bonds over more than one species implies that the displays are homolo-
many years. It and similar species are said to 'mate for life'. gous; that is, evolutionarily derived from a single display of
In most cases, they re-mate as long as they both return to a common ancestor. Certain displays show close similarities
their previous site at the right time. If not, new mates may among the ibises and storks, and the herons (Hancock and
be taken. Several species of tropical ibises and several Kushlan 1984). However, because of uncertainty as to the
storks, such as the Saddlebill, Blacknecked and Jabiru, are phylogenetic closeness of these groups, it is best to maintain
solitary, non-migratory, and may remain together through- a separate behavioural nomenclature, unless the similarities
out the year. These pairs may not require extensive court- of the displays are so compelling as to lead us to imply their
ship at the start of the next breeding season. homology.
Pairs form during a courtship period in which both birds In general, the process of courtship starts when birds
engage in a series of behaviours that help establish and then begin to gather near a potential nesting site during the
maintain the bond. The initial stages are crucial in that the daytime. Depending on the species, these sites may be
birds have to overcome the tendency to exclude other birds patches of trees, marsh reeds, or cliffs. In most cases the
nearby. That is, for much of the year, a bird tries to keep nesting sites are characterized by some degree of remoteness
other birds at a comfortable distance, yet, when pairing, is or inaccessibility, and many are located on islands. In these
faced with accepting the close proximity of another bird.
One correlate of pair formation in most species is a
seasonal change in the colour of the bill, face, legs or other
exposed skin, called the soft parts. In the American White
Ibis, the face turns a bright red and the bill black, and the
females develop a red gular pouch under the chin. In the
Sacred Ibis, the skin along the breast and under the wings
turns bright scarlet and is highly visible when the bird flaps
its wings. The Sacred Ibis also develops ornamental plumes
at this time. The Oriental Crested Ibis has a unique type of
cosmetic coloration. It uses its head to paint its pinkish back
feathers grey with a secretion from its preen gland. These
colour changes enhance the displays of the birds.
The overall process is more similar between species of
storks, ibises, and spoonbills than it is different. However,
the details of the behaviours used in the process, called
displays, differ among species. Species-specific courtship
displays reduce the chances of mating with the wrong
species. The occurrence of various displays and the different
forms of display can serve as distinguishing characteristics
between the species, just as morphological and plumage
differences do.
Most of the displays shown by storks, ibises and spoon-
bills during their courtship are 'derived' activities (Tinber-
gen 1952). These are behaviour patterns —usually borrowed
from one or more other activities such as preening, fighting
or fleeing™that have evolved as social signals. Displays
have become specialized through a process known as 'ritua-
lization' (Blest 1961), in which the elements of the display
have become elaborated and stereotyped into the species-
specific patterns that are recognized by the recipient bird.
The various displays used by storks, ibises and spoonbills
can be defined and categorized. Many of these behaviours
have been previously described and named (Kahl 1966a, Figure 23. American Wood Stork. Social interactions be-
1971f, 1972a-d, 1973b, 1983, 1988a). Displays are named tween birds in a treetop colony (southeastern USA).
22 Courtship and Reproduction
Figure 24. A Yellowbilled Stork feeding by tactolocation
with bill held open and moved slowly about in water (cen-
tral Kenya).
Figure 25. A Painted Stork sunning in the 'delta-wing'
posture (northern India).
locations males behave in ways that serve to attract females
and to advertise their readiness to mate.
Most information available is on colonial species, and
generalizations are often based on these species. It can be
expected that solitary species may have slightly different
approaches to mate attraction. In some non-colonial
species, individuals remain on an activity area year-round
and may return to the same nest-site in succeeding years.
The Hadada Ibis appears to exhibit pair-bonding behaviour
even within a flock, as pairs of birds separate out to preen
mutually.
In colonial species, the male birds stand in groups in a
potential colony site and periodically fly about. The colony
forms as displaying males and interested females gravitate
toward each other, forming 'bachelor parties'. It is usual to
find that a large colony is composed of smaller groups in
which all the pairs are in the same stage of nesting, as a
result of displaying birds having been attracted to each
other.
A male eventually selects a display perch within the
selected colony site in a location where he is visible to
interested females. In most species the eventual nest-site is
located at or near the display perch, so it is possible that the
quality of the site a male is able to secure and hold against
competing males may be of considerable importance in
formation of the pair. Mate choice is primarily a matter of
the female selecting from among available, displaying
males.
The displays of the male also serve to intimidate nearby
competing males. When the defensive content of a display is
insufficient, fights can develop among males. The female
approaching its chosen male also has to overcome his
aggressive tendencies, and some of her displays at this time
can be interpreted as appeasements. In the American White
Ibis, the much smaller female can become quite bloodied
during the process of gaining access to the male's defended
display site.
After mate selection is made, the pairing is reinforced by
additional displays that often continue for several days.
These eventually lead to copulation. Since the displays
leading up to pair formation and copulation are defined
differently for the two groups (storks, and spoonbills and
ibises), it is worthwhile to describe these separately.
COURTSHIP IN STORKS
In all species, courtship begins with a male making himself
conspicuous and available to an interested female. In many
situations, the male just stands conspicuously. During
Standing, a bird may try to look imposing. The Marabou
blows up its gular air sac.
One of the first active advertising displays used by male
storks of the genus Mycteria in their bachelor parties or on
their display sites is Display Preening, in which the bird
repeatedly goes through the motions of running its bill down
the primaries (the main flight feathers) of one wing then the
other. It does this just as if preening, but the feather itself is
not grasped, and perhaps not touched at all.
Often occurring during episodes of Display Preening,
 Courtship and Reproduction 23

Swaying Twig-Grasping display is executed by the male

bending forward to one side and grasping a twig. Upon
releasing the twig, the bird sways to the other side and
repeats the process. A similar advertising behaviour found
only in the openbills is the Advertising Sway. With bill
gaping, the bird bends its head to its feet and lifts its wings
slightly away from its body, while shifting its weight from
foot to foot and swinging its bill from side to side.
The Head-shaking Crouch, is given by storks of the genus
Ciconia^ when a female approaches a displaying male. He
inclines his body forward with head and bill low, wings
slightly away from body and, while shifting from foot to foot,
shakes his head in a 'no' motion, sometimes lowering the
body to the nest in the process.
A ritualized response to disturbance is the Arching Dis-
play, given by all stork species. In this 'anxiety' display, the
bird, from an erect posture with feathers sleeked down,
leans forward and looks at the offending object. In some
species this may be accompanied by spread wings and
vocalization.
The generalized agonistic display among storks is the
Forward Display. With the neck retracted, body horizontal,
and feathers erect, the bird points its bill at its opponent and
moves toward him. An antagonistic bird may also rattle its
bill in a Forward Clattering Display. Although ritualized,
these threat displays can lead to overt fighting should the
intruder not choose to retreat. A hostile bird may pursue an
opponent into the air using an Aerial Clattering Display.
The pursuing bird flies after and above the opponent with
its neck extended and bill pointed toward him. The attack-
ing bird may clatter its bill but usually does not try to bite
the opponent, who usually takes evasive action.
An intriguing, apparently agonistic, display is called
Mock Fighting. On or near the nest-site, a bird suddenly
erects its neck feathers and makes lunging grabs into the air
as if fighting or catching a flying insect. The bird frequently
loses its balance and has to flap its wings to stay on the
perch.
The usual subordinate display of storks, is called the
Upright Display. The submissive bird stands erect with
head extended upwards, feathers compressed, and turns
away from the threatening bird. At least two appeasement
displays are used by the female stork in attempting to gain
access to the male's courtship station, Balancing Posture
and Gaping. Upon approaching the male, the female (of
some species) assumes the Balancing Posture, holding her Figure 26. Asian Openbill Stork. The Up-Down Display
head low with her wings spread. This may be accompanied at a nest with large young (south-central Thailand).
by bill Gaping, as if overheated, but the bird does not
actually pant. As time passes without being driven away, bill. The display is used when levels of hostility are still high
she may close her wings but often continues to Gape with among the newly paired birds. A stork may infrequently
her head bowed. give a bill snap when taking flight. During this display, the
Soon after acceptance of a mate, a male Mycteria may Pre-flight Snap, the bird bends forward and low, swinging
engage in a Flying Around Display. He takes off from the its head between the legs. It then gives a single loud snap
nest and flies in a tight circle before returning to land next to with its bill followed by a normal take off. Another display
his mate. given before flight is the Erect Gape. The bird stands erect,
Also during or soon after pair-formation, Bill Snaps be- with neck extended almost vertically and bill held slightly
come common. This behaviour, shown by both sexes in below the horizontal. Sometimes a low call is given. The
some species, involves a quick lift of the head to bring the Erect Gape may also be followed by a Bill Snap.
bill to slightly above horizontal, followed by a snap of the The Flap-Dash display —probably derived from foraging
24 Courtship and Reproduction
Figure 27. A female Greater Adjutant Stork preening
near her nest (northeastern India).
behaviours —is performed on the ground or in shallow water
by the Saddlebill, Blacknecked and Jabiru Storks. A bird,
usually the male, flapping its wings, runs several metres
away from its companion and then runs back again.
Following pair-formation, mates greet each other with an
Up-Down Display, which is performed by all storks but
differs among species. In the openbills and wood storks, the
bird lifts its head vertically, and, with its bill gaping, it may
utter short hissing or grunting vocalizations, while lowering
the bill toward the floor of the nest-platform. Depending on
species, during the head-up part of the display, the bird may
snap the bill once or twice or repeatedly, or it may not snap
its bill at all. In the Blacknecked Stork, the bill is not raised,
but the wings are spread and fluttered and the bill is
clattered rapidly. Other species show varying degrees of
head movements and clattering during the Up-Down. The
most extreme is reached in the White and Oriental White
Storks, in which the head is thrown all the way back toward
the tail and the bill-clattering is loud and prolonged. This is,
perhaps, one of the most widely known of all stork behav-
iours. Up-Down Displays continue throughout the nesting
season.
Copulation occurs at or near the nest-site with no pre-
copulatory display. Characteristic of storks is for the male to
clatter its bill or rattle it against that of the female during
copulation. In storks, this behaviour is called Copulation
Clattering.
A Nest-Covering Display has been observed in several
species. The parent bird hides its eggs or young under
partially opened wings.
COURTSHIP IN IBISES AND
SPOONBILLS
The courtship displays are more poorly known among ibises
and spoonbills than in storks. Many species have never been
studied at all. None the less, from the information available,
a number of displays can be identified. Certainly, additional
studies on these species are needed.
As in storks, the males of colonial ibis and spoonbill
species, and those that must pair each year, display for
females. Upon choosing a display site, ibises and spoonbills
tend to remain on it almost continuously. In fact, Display
Standing may be considered the initial display in courtship.
Display Standing can develop into Display Sleeping. In
imitation of sleeping, the bird places its bill under its back
feathers, although its eyes remain open. This behaviour
appears to be quite contagious among neighbouring birds.
It also occurs as a pause in active display, serving no doubt
to maintain a bird's continued claim to a display- or nest-
site.
Species that remain paired throughout the year perform
displays that serve to reinforce the pair bond, and perhaps
to coordinate breeding readiness. Pairs display together
with simple behaviours such as Standing, Head-Shaking,
Stick-Shaking and allopreening.
Ibises can be quite noisy when nesting, even those that
are rather silent for most of the year. Calls differ among
species from whistles to caws; they also may make noise
with their wings when landing, taking off or flying around.
Ibises and spoonbills use calls for contact, to communicate
with mates, during copulation, and in agonistic situations.
Some species, such as the Barefaced Ibis, perform duets
(Luthin 1983a).
The active solicitation behaviour of unmated males, in
species so far studied, consists primarily of three displays:
Display Preening (Front and Rear), Display Shaking, and
Bill Popping. Display Preen (Front) and Display Preen
(Rear) occur when the bird appears to use its bill-tip to
preen its primary feathers (Front) or to preen over and
behind the wings (Rear). The Display Shake is given by a
bird standing erect and holding its wings loosely. It then
shakes the wings alternately up and down. The Bill Popping
display occurs as the bird snaps its gaped bill shut, usually
without sound, although at times accompanied by a quiet
pop. In African Spoonbills, this display is given as the bill is
raised to the horizontal or slightly above. In the ibises so far
studied, it is given with the head extended and bill down-
ward. Ibises sometimes also grasp a twig and shake it
briefly. Bill Popping is very reminiscent of the Snap Display
of herons and also is reminiscent of the Bill Snap of storks.
The solicitation displays may be given in any direction,
not necessarily in the direction of a female, and sometimes
toward neighbouring pairs. In some ibises, Bill Popping
may be truncated into a more moderate form in which the
neck is not extended. In ibises so far studied, the three

displays are often used together but not in any standard
sequence. In the African, Royal and Yellowbilled Spoon-
bills, these three displays are given in succession: Display
Shake/Display Preen/Bill Popping.
Another display, observed primarily in courting female
spoonbills, is the Display Flight. The bird takes to the air,
flying in a low circle or even a figure-of-eight while calling.
It then lands near its point of origin, which is usually near
an unmated male. This display is similar to the Flying
Around of the storks.
The initial phase of pair-formation involves considerable
aggressiveness by the male toward the intruding female. As
a result, many of the initial behaviour patterns of courting
male ibises and spoonbills have components of aggression or
fear. When a bird is alarmed, the Erect Posture is assumed.
The bird stands upright, feathers sleeked, bill down, peering
at the disturbing object. This behaviour is similar to the
Arching display of storks and may precede, but does not
necessarily lead to, escape behaviour.
The basic aggressive behaviour of ibises and spoonbills is
the Sparring Display, in which the bird assumes a horizon-
tal posture and moves toward or lunges at an intruder using
a bill thrust. The Sparring bill thrust may or may not make
contact. An overt fight may follow. Two birds engaged in
ritualized Sparring hold their feathers erect and bills gaped,
Figure 28. Asian Openbill Storks. High water levels pre-
vented breeding at Bharatpur, India.
Courtship and Reproduction 25
while each reaches forward and snaps its bill towards the
antagonist; it does not actually make contact. This behav-
iour is returned by the opponent, also without contact, and
a stab-and-counterstab Sparring match commences, with
birds alternating ritualized attacks. The aerial modification
of Sparring is the Supplanting display, in which a bird flies
at another bird, with its neck extended and bill gaped. The
attacker slows its flight with its wings, and attempts to land
on the opponent or, more likely, on the perch vacated by the
displaced victim.
One of the most common activities of some species of
ibises is appearing to rub their head on their back. The
Head Rub is ritualized into a mutual appeasement display
and also an advertising display. The bird places the side of
its head on its shoulders, bill pointing upwards. The bird
may then turn it slightly to vertical or even to the horizontal
on the other side, in the rubbing motion, or it may make
only a quick touch of the nape to the upper back. The female
ibis uses this display prior to attempting to enter the male's
display area, and the male may use it with other attraction
displays. The origin of this display is clearly the similar
comfort movement. In some species it is reminiscent of
aspects of the Up-Down display of storks. In its vertical
form, it is similar to the Stretch Display of herons.
Upon actually attempting to enter the male's display
area, the female assumes a Bowing Display, in which she
approaches the male keeping her body and head low,
feathers sleeked back. She turns her head away from the
26 Courtship and Reproduction

male, exposing the side of her head to him. The male

generally reponds with a Sparring display and even an overt
attack, in response to which the female may flee but does not
fight back. The male responds initially by aggressive actions
and eventually by acceptance of the female at the nest-site.
This behaviour is identical in function and similar in
appearance to the stork's Balancing Posture, except that the
wings are not opened.
The aggressive phase of pair-formation ends as the male
accepts a female. Bowing Displays continue, and when one
of the pair returns to the nest, a Greeting display is given.
With head feathers erect (if present) and bill gaped up-
wards, the bird calls and may flap its wings. In spoonbills
the Greeting is given with the bill upward, whereas in ibises
the head and bill are slowly brought downward to the
vertical in a bowing motion, before the vocalization. Espe-
cially in spoonbills the downward component is recogniz-
able as a distinct display, Head Shaking. Bending forward
simultaneously with bills downward and closed, the birds
shake their heads in a 'no' motion while lowering them to
the nest. Some ibises also Head Shake. The birds' necks
may overlap, the taller male usually being on the top. The
Greeting displays appear similar to the Up-Down of storks.
Stick Shaking may also occur. Bending forward with bills
pointed down, the birds grasp nest material and shake their
heads rapidly from side to side. Again, necks may cross and
overlap, with the male on top. This behaviour is similar to
Head Shaking, except for the involvement of twigs.
Another mutual display of paired birds is the Head
Quiver. Standing upright with head feathers erect, a bird
twitches its head several times. Generally the display is
performed by both sexes in succession or simultaneously.
Following the acceptance of a mate, pair-formation is
completed through copulation. Allopreening continues as do
Head Shaking and other Greetings. During the Copulation
Display, the male climbs on the back of the nearly horizontal
female, and brings his cloaca into contact with hers, while
flapping his wings for balance. In the process, the male may
also perform a bill-shaking behaviour in which he grasps the
female's head or bill or holds his bill alongside her neck while
he shakes his head or nibbles at her feathers.
A later defensive posture is the Nest-covering Display. A
parent bird erects its feathers, orients toward the threat, and
droops its wings down and over the nest. This display is
clearly homologous to the behaviour of the same name in
storks.
Birds of a pair often engage in preening each other (allo-
preening). They especially nibble the feathers of the
partners head and neck. Throughout nesting the birds com-
monly occupy themselves with preening, head-rubbing and
shaking. Although these are functional maintenance behav-
iours, they probably also serve a display purpose, helping to
maintain the pair-bond.

Figure 29. A Eurasian Spoonbill with well grown young

(Bharatpur, India).

Figure 30. Young Asian Openbill Storks leave the nest

early (Bharatpur, India),
 Courtship and Reproduction 27

NESTING
After courtship and pair-formation, nesting begins. Depend-
ing on the species and the habitat, nests may be made out of
sticks, reeds or vines, often with green material added. Nest-
building is generally the province of both sexes. The male is
largely responsible for gathering the sticks. The female
accepts the material. The sticks are then inserted by the
female into the developing nest bowl by a behaviour called
'tremble-shoving', holding the stick by an end and wiggling
it into the nest matrix. Behaviours during nest-building and
maintenance may involve the real or feigned picking, grasp-
ing or presenting twigs to the female.
At this time, appeasement and pair-formation displays
become less frequent and less intense. The eggs are laid a
day or more apart, and incubation usually begins with the
first or second egg. The members of the pair alternate
incubation duties. The larger male is often in attendance at
the nest for much of the day.
Young hatch asynchronously and are unable to stand
from the first ('altricial young'). The bills of hatchlings are
short, straight and rounded, even in ibises and spoonbills.
The typical shapes begin to develop soon thereafter.
Young are fed by both adults on regurgitated food. Storks
regurgitate food onto the floor of the nest and their nestlings
pick it up, whereas nestling ibises and spoonbills insert their
entire head and bill down the parent's gullet to receive food.
During the first few weeks after hatching, one adult remains
at the nest at all times, as the parents alternate foraging
trips. Storks also bring water to nestlings.
When the young are further developed, both adults leave
the nest to forage at the same time. Nestlings respond to
disturbance with threat and alarm displays. Storks respond
with a Nestling Bristle and may give Forward Displays.
Ibises and spoonbills assume an Erect Posture.
Nestlings solicit food from their parents using a Begging
Display. Facing the parent while resting on its tarsi, the
nestling raises its head and bill, bobs them up and down,
flaps its wings, and gives a begging call. As the nestlings
grow they may give the display while standing or—
especially in the case of the smaller, more active ibises and
spoonbills—while running or even flying after the parent.
Larger young may catch their parent, and placing a wing
over the parent's shoulder, pin him or her down while they
force feed from the gullet.
Large young spend considerable time flapping their
wings. In some colonial species, it appears that young leave
their nests to form groups, called creches. The young can fly
long before they depart from the colony. They usually re-
turn to near their nests to be fed by the parents, which they
appear to recognize by call and sight. Fledging periods vary
widely, according to the size of the species, from less than a

Figure 31. Storm's Stork. A unique photo of an adult

standing on an active nest; only two nests of this species
have ever been found (in Thailand and Sumatra). Photo by
courtesy of the late Seub Nakhasathien.

Figure 32. An Asian Openbill Stork in Bharatpur, India.

28 Courtship and Reproduction

month in the smaller ibises to more than 3 months in the

larger storks.
Although for most species, parental care ends soon after
fledging, the young of some species of ibises —those that
appear to maintain long-term pair-bonds — continue to use
the family's home range for a time, and are even fed there by
the parents.

Changes to display names

Original name New name

Storks1 Forward Threat Forward Display

Aerial Clattering Threat Aerial Clattering
Forward Clattering Threat Forward Clattering
Snap Display Bill Snap
Anxiety Stretch Arching Display
Nestling Defence Display Nestling Bristle
Spoonbills and Alarm Posture Erect Posture
ibises2 Appeasement Display Bowing Display
Flight Intention Pre-flight Tipping
1
Used in Kahl 1966a, 1972a-d, 1973b.
2
UsedinKahl 1983, 1988a.

Figure 33. A captive Oriental White Stork at Vogel Park,

Walsrode, Germany.

Figure 34. African Spoonbills Sparring and nest building

on a rocky island on Lake Turkana, Kenya.
 Feeding Behaviour
and Ecology

s
torks, ibises and spoonbills are among the more typical of the 'wading' birds, species that
characteristically feed while walking about in shallow water. How these birds behave while
feeding and what they can catch to eat are constrained by a general adaptive plan, millions of
years old. That design includes long legs, long necks and long bills, all adaptations for
feeding while walking through shallow water.
Because of this body plan, most species of storks, ibises, and spoonbills are highly aquatic, character-
istically occurring in and around shallow water. The wood storks and the openbill storks epitomize a
near total dependency on water and are found only in very specific types of aquatic habitats. However,
most species can also feed on land should compelling suitable food become available there. Some, such
as Abdim's Stork and the Waldrapp, Bald and Hadada Ibises, are more terrestrial than aquatic. They
feed mostly on insects in dry grasslands and farm fields. Yet they retain the primordial body design,
modified by somewhat shorter legs, necks and bills.
All species are both liberated by and constrained by their wading bird heritage. Indeed, it is
instructive in understanding the ecology of these species to consider what foraging opportunities can be
possible for large birds built according to these general specifications.
The various behaviours used by storks, ibises, and spoonbills in their foraging activities can be
delineated and defined (Kushlan 1978a). Recognizable similarities in most behaviours among the
species allow use of the same terminology in both families. In this book we capitalize the named feeding
behaviours in order to allow the reader to identify them in text. We also capitalize, as is the custom,
courtship behaviours. However, unlike courtship behaviours, most feeding activities are not ritualized
displays.
The number of behaviours identified as being used in feeding is surprisingly extensive. No species is
known to feed in all the ways so far documented. Some species do exhibit quite a variety. The Roseate
Spoonbill, for example, has been documented as being able to use eight different feeding techniques.
The most universal feeding behaviour is Standing. That is, a bird simply stands in one place and
captures prey as it is encountered. Standing can be used on land or in the water, or even while mobile—
African Openbill Storks Stand on the backs of moving hippos (Kahl 1971c). Typically, Standing
foraging involves capturing prey that is on the ground or in the water. Two special types of Standing
behaviour have been recognized when prey occur other than in the water. When Standing Flycatching,
a stationary bird catches prey that is airborne. When Gleaning, a Standing bird catches prey that is on
an emergent plant or some other above-water object. Another variation, which occurs while Standing, is
Flipping. A bird turns over an object to catch what is underneath. In Foot Stirring, a bird vibrates its
foot or leg, to stir up potential prey.
No species, or even an individual, forages solely by Standing, because prey-capturing opportunities
are soon depleted at most feeding sites. To move from place to place, birds intersperse Standing with
more mobile behaviours. The most widespread behaviour is Walking Slowly, in which a bird walks or
wades at slow speed usually continuing to seek prey. Faster passage is achieved by Walking Quickly
and by Running. Walking Slowly can be recognized by the bird's slow rate of progress, as slow as a step
or two per second. Running, on the other hand, can be recognized by its rapid rate of progress, such that
the bird occasionally needs to use its wings for balance. Of course, these classifications are relative and
not totally distinctive.
30 Feeding Behaviour and Ecology
The relative amount of Standing and Walking depends on
the species and the situation. The Shoebill typically Stands
in one place for long periods at a time, whereas ibises and
spoonbills seem always on the move.
The usual prey-capturing technique involves a Bill Grab
in which the bill grasps a prey item, using the mandibles as
pincers. Unlike herons, which strike from a recoiled neck
(Hancock and Kushlan 1984), storks and ibises Grab prey
without a darting stroke. The Shoebill is unusual in that its
Grab is accompanied by a Body Lunge, in which the head
and trunk are thrown at the prey. The bird often ends up
partially submerged (Guillet 1979). Some food can be cap-
tured in a more leisurely manner by Pecking. A bird simply
uses its bill to pick up an item from the water or substrate.
Wood storks use a highly specialized prey capturing behav-
iour, the Bill-snap Reflex. Upon contact between a prey
item and the open bill, the two mandibles close refiexively at
a speed averaging 25 ms in the American Wood Stork (Kahl
and Peacock 1963).
Several special techniques have developed to allow birds
to forage without using their eyes. The most widespread is
Probing, in which a bird quickly and repeatedly moves its
bill tip into and out of the water or substrate. This is the
characteristic non-visual foraging method of ibises, which
have innervated bill tips that register contact with a prey
item. It is also used by many storks, including openbills,
typical storks and giant storks. In ibises several types of
Probing have been distinguished according to water depth
and rapidity of the probe (Kushlan 1977a). One can recog-
nize Shallow Probing (the bill is inserted fewer than 2 cm
Figure 35. Immature Painted Stork (Bharatpur, India).
into the substrate), Deep Probing, Standing Probing, Walk-
ing Probing and Multi-probing (Walking interrupted by
Standing Probing). Groping is a similar behaviour in which
a bird places and moves its open bill about in the water. It is
the characteristic component of the feeding repertoire of the
wood storks.
Head Sweeping (Head Swinging of Kushlan 1978a; Slow
Sweeping of Vestjens 1975b) is the typical tactile feeding
behaviour of spoonbills but is also used by ibises. Vestjens
(1975b) showed that the details of Head Sweeping differed
in the two Australian spoonbills. It is usually accomplished
while Walking Slowly. When performed while standing still
and searching a site with short sweeps, it is called Intensive
Sweeping. Forward Ploughing takes place when a bird
walks with its bill under water and projecting forward,
rather than swinging it sideways. Dragging is similar except
that the bill is dragged through the water at the bird's side.
Wings are also used during foraging. Feeding while actu-
ally in flight has not been recorded in storks, ibises or
spoonbills, although it can be common in some species of
herons. Feeding birds do take short repositioning flights,
however. In Hopping, a bird flies up and alights nearby,
even adjacent to its former feeding spot. In Wing Flicking, a
bird quickly partially extends and retracts its wing; whereas
in Openwing Feeding, a bird completely extends and re-
tracts one wing at a time. These behaviours appear to serve
to frighten prey, especially fishes, into movement.
Several behaviours tend to be strung together in various
ways to create a feeding sequence. The wood storks, for
example, typically forage by alternating Standing with
Figure 36. An Oriental White Ibis at Bharatpur, India.

Figure 37. African Openbill Storks feeding on snails in a lily pond (Tana River, Kenya).
Walking Slowly, while Groping with their open bill and
Foot Stirring. At the same time they Wing Flick or Open-
wing Feed on the same side as Foot Stirring. The mix of
these actions increase the activity of potential prey, and
therefore the chances of its swimming into the bird's Grop-
ing bill (Kahl 1964, 1972b).
The Blacknecked and Saddlebill storks Walk Slowly or
Run while Probing vertically in the water with each step.
The American White Ibis is typical of a bird with less
restricted sequencing (Kushlan 1977a). It will Walk Slowly
while Probing; then, stopping at the edge of the water it may
Peck at the mud surface or attempt to excavate a crayfish
from its burrow by Standing Deep Probing. It then might
move to deep water where it will Head Swing like a spoon-
bill.
Once a bird captures a prey item, it must be 'handled'.
Small prey can be consumed immediately. Ibises can swal-
low such items nearly instantaneously by closing their bill
tips while slightly pushing their head forward. Large prey
have to be softened or broken into smaller pieces. A large
food item, dead or alive, can be handled by Biting (i.e.
mandibulating the prey) or by Stabbing (i.e. repeatedly
poking and piercing it with the bill). The Australian White
Ibis holds mussels against a rock with its feet whilst stab-
bing them with its bilL Handling time goes up exponentially
Feeding Behaviour and Ecology 31
as prey size increases, soon reaching impractical limits such
that a bird might forgo eating a captured prey item in favour
of resuming the search for a new one (Kushlan 1979c).
Divergence among the several lineages of storks, ibises
and spoonbills is reflected, indeed in large part influenced,
by their feeding behaviours and accompanying morphologi-
cal features. One major evolutionary and ecological division
is among species that feed primarily visually, by seeing their
prospective prey before catching it, and species that feed
primarily tactilely, by feeling prospective prey with their bill
(Kushlan 1978a). It has been shown that the American
Wood Stork feeds just as effectively without use of its eyes as
with them (Kahl 1964). The primarily visual foragers, in-
cluding the typical storks and adjutants, generally Grab the
prey items they see. The primarily tactile foragers, including
the ibises, wood storks, openbills and spoonbills, typically
use Probing, Groping and Head Sweeping. Despite these
overall adaptive trends, most species have the flexibility to
use both visual and tactile methods when the appropriate
occasion presents itself.
Given the correlations between feeding behaviour and
morphology, much of the feeding story of each species can
be told by examining their primary feeding apparatus, their
bill. The divergence between ibises and spoonbills is essen-
tially one of billshape. The ibises have thin downturned bills
 32 Feeding Behaviour and Ecology

whereas the spoonbills have straighter, flattened bills, each

representing a basic morphological type for tactile aquatic
foraging. Ibises that feed in terrestrial habitats, such as the
Green Ibis, have shorter bills than those that feed in water,
such as the American White Ibis.
The storks have an even greater diversity of billshapes.
Ranging from the enormous Grabbing bill of the Jabiru
Stork, to the specialized bills of the openbill storks and the
ShoebilL Openbill storks probe for snails, capturing and
opening them in their bill-tips. The Shoebill feeds visually,
Grabbing its very large prey. The wood storks have slightly
down turned bills, reminiscent of the ibises.
Another peculiar morphological correlation with feeding
behaviour is the colour of the feet of the American Wood
Stork. Its feet are pinkish, contrasting with its darkish legs.
This colour possibly serves to enhance the effectiveness of
Foot Stirring while feeding.
Some species are solitary foragers. They often use simple
feeding behaviours such as Standing or Walking Slowly,
stalking prey in ways that are not enhanced by a crowd. The
Shoebill is the archetypical example of a solitary species.
Others are solitary at one time of year and more social at
another. The Spotbreasted and Hadada Ibises are two
examples of species that nest solitarily but feed in groups
when not breeding.
The great majority of species are capable of feeding in the
company of other birds. Generally speaking most storks,
ibises and spoonbills feed at least some of the time in social
associations, called feeding aggregations, which form at sites
of high food availability. Others show the higher level social
organization typical of flocking.
A correlation between nesting and feeding dispersion is
too great to be a coincidence. Colonial nesting and commu-
nal roosting provide exceptional opportunities for feeding
together (Krebs 1978). Birds nesting in the same place can
take advantage of foraging opportunities found by other
birds in a process called social facilitation. Either by follow-
ing birds flying to the feeding grounds or by seeing other
birds feeding, wading birds are able to identify promising
feeding sites and join the aggregation of foraging birds
assembled there.
Many of the communally foraging species are white, a
colour that has been demonstrated to attract wading birds
to a feeding site (Kushlan 1977e). This raises an interesting
question: once a bird finds a profitable foraging site, why
should it advertise the place to other birds? At least two
non-exclusive possibilities present themselves. The first re-
lates to the character of the feeding sites used by most of
these species. These sites, such as drying ponds and shallow
wetlands, are ephemeral in that they are suitable for feeding
only for short periods. Also in such cases the amount of food
present is more than one bird can eat in the short time it will
be available. Thus there may be no harm in inviting other
birds to join the feast. The second possibility is that a
foraging bird actually benefits from having more birds at its
Figure 38. A Hadada Ibis feeding in Kenya. feeding site. One benefit may be in hiding from predators
within a large aggregation of other birds. More directly, the
Figure 39. A Buff-necked Ibis in Argentina, South prey present are disturbed by the feeding activity, which
America. increases the probability of one bumping into a waiting bill.

This is a form of mutualism, a form of interaction between
individuals or species from which both benefit. A number of
instances of commensalism (an interaction in which one
animal benefits and the other is unaffected) have also been
reported in storks, ibises and spoonbills. Generally a
visually foraging species follows a tactilely foraging species,
benefiting from the prey the latter stirs up (Kushlan 1978a).
The subtle interactions among species, as noted above,
may have lead to the social signalling function of the white
plumage found in many species of storks, ibises and spoon-
bills (Kushlan 1978a).
Feeding aggregations also include other aquatic birds,
especially herons and cormorants, though aggregative for-
aging carries with it certain risks, especially that of losing a
captured prey item to a robber. Small birds can fly over and
snatch a prey item from the bill of an ibis or stork. Larger
wading birds also steal from smaller wading birds (Kushian
1978c, Kushlan et al. 1985). Especially at risk are prey that
take time for handling.
Some species that feed in family groups may tend to feed
in more permanently available feeding sites. Species that
remain in family groups, represented by the Blacknecked
and Saddlebill storks and several species of ibises, also use
the same foraging area for large portions of the year. Mem-
bers of the pair forage within sight or hearing of one another
but only join aggregations if one happens to form in their
home range.
Most storks, ibises and spoonbills feed primarily during
Figure 40. White Storks feeding on drained carp ponds in Israel.
Feeding Behaviour and Ecology 33
daylight. However, some — especially among the tactile
feeders —will forage at night, and even visual feeders can
secure prey on bright moonlit nights. In general, the smaller
species begin feeding at sunrise. Storks often do not leave
the colony site for feeding grounds until later in the day.
This is because they travel long distances, not by flapping
flight, but by soaring high on thermals and then gliding a
long way to the next thermal. Such species leave the colony
or roost when the ground heats up in mid-morning, forming
the thermals. In certain species, such as the American
Wood Stork, some individuals may spend the night at the
foraging ground, returning to the colony the next morning
(Kahl 1964). Outside the nesting season, storks, ibises and
spoonbills spend much of the middle of the day resting near
the feeding ground.
Storks, ibises and spoonbills are overwhelmingly car-
nivorous, each species eating various sorts of fish and am-
phibians, snails, crustaceans, insects and other aquatic in-
vertebrates. The bulk of the diet of a population may consist
of very few species of prey.
The most specialized group is the openbill storks, whose
peculiar bill allows them to extract aquatic snails from their
shells (Kahl 1971c). Thus snails make up almost all of their
diet. With their Probing bill, ibises are especially adept at
capturing small invertebrates, such as prawns, crayfish and
aquatic insect larvae that live on the bottom or on aquatic
plants. Similarly, they can probe into loose soil or grass to
extract terrestrial insects and their larva. Faster-swimming
34 Feeding Behaviour and Ecology
prey, such as fish, are also caught but usually only con-
sumed in numbers when they are available in high densities.
With their Head Sweeping, spoonbills take prey from the
water column or in submersed plants, generally catching
fish, insects and prawns. The wood storks are restricted in
diet by their tactile foraging techniques. Their food is pri-
marily fish that are found in rather dense concentrations
(Kahl 1964, Kushlan 1979b).
Several groups depend heavily on scavenging for food for
most of the year. The adjutants are specialized scavengers
on large prey killed by predators, and the Sacred Ibis often
scavenges at kills and dump sites. Scavengers revert largely
to living food when feeding their nestlings. The typical
storks have the most generalized diet, being able to select
their prey visually. In addition to the expected fish and
amphibians, they also consume small mammals, insects and
reptiles, depending on what is available.
Prey availability is, in fact, a key to understanding the
foraging ecology of these species (Kahl 1964, Kushlan
1978a, 1981, 1986b). Many of the fundamental adaptations
of storks, ibises and spoonbills are related to accommodat-
ing to changing patterns of prey availability. These include
seasonal migration, intraregional population shifts, timing
of the breeding season, colony site selection, foraging habi-
tat use, and food consumption.
Those species that have relatively specialized bill shapes
and feeding behaviours often depend on very specific cir-
cumstances of prey availability, generally related to
seasonal patterns of rainfall and water level fluctuations
(Kushlan 1978a, 1981, 1989a). Aquatic ibises require shal-
low water; terrestrial ibises require ground conditions suit-
able for probing. Wood storks require high prey densities for
efficient foraging, and their nesting success depends on the
continuous availability for several months of shallow, ephe-
meral foraging sites. The less specialized typical storks can.
forage in either wetlands or fields and depend on seasonal
changes in food availability.
The migratory patterns of many species are correlated
with changing food supplies. The White Stork depends on
frogs during the nesting season in Europe and insects —
Photographs. The photographs appearing in the above sections were kindly supplied by M. P. Kahl (Figures 1-27),
J. A. Hancock (Figures 28-30, 32-40) and the late Seub Nakhasathien (Figure 31).
especially plague locusts—in the winter in Africa. The sur-
vival of storks during the non-breeding season has been
shown to depend on rainfall patterns and therefore food
availability on the African wintering grounds.
Fires can increase the prey available to upland foraging
birds. Several species, such as the Marabou, Maguari,
White and Woollynecked Storks and the Sacred and Bald
Ibises, characteristically forage on recently burned land,
even following the fires closely to catch insects and other
prey exposed by the fire.
Prey availability is especially critical for the tactile for-
agers (Kahl 1964, Kushlan 1979b,c). Their ultimate diet is
the result of the interaction of their size, bill shape, feeding
behaviour, sizes and types of prey available, and prey abun-
dance. As a result the American White Ibis and American
Wood Stork, feeding in the same area, take entirely different
diets, owing to differences in where they can feed and what
they can catch and handle.
Food is required for the maintenance of adults and for the
raising of young; the latter can as much as double the daily
energy needs of a pair of adult wading birds. All the storks,
ibises and spoonbills are relatively large birds and have
substantial daily energy requirements for growth and main-
tenance (Kahl 1962, 1964, 1966a, Kushlan 1977b,c, 1978a).
The daily energy needs of an American White Ibis has been
calculated to be about 200 g of food, whereas the larger
Marabou Stork requires 720 g. Nestlings increase this
requirement.
The rigours of delivering sufficient food to nestlings deter-
mines much about the seasonality of breeding and feeding
habitat selection exercised by these birds. Nesting failure is
not uncommon, generally brought about by changes in
rainfall patterns, which affect food availability. Because the
adults begin incubating their eggs as soon as the first is laid
rather than waiting for the entire clutch, the young hatch at
different times, i.e. asynchronously. As a result the older
chick gets a head start on its siblings and may survive food
shortages when the younger birds perish, thus permitting
some production under difficult conditions.
 American
Wood Stork
Mycteria americana Linnaeus

Mycteria americana Linnaeus, 1758, Syst. Nat., ed. 10, p. 140: Brazil

Other names:
Wood Ibis, Flinthead, Ironhead, Gourdhead, Gannet, Preacher, Spanish Buzzard, Colorado Turkey,
Wood-Pelican (English); Gaban, Gaban heucito, Garzon, Coco, Cabeca seca, Cigiienon, Doroteo,
Sargento, Ibis del Monte, Alcatraz, Sowewies, Negroscopes, Cabeza de piedra, Galletan, Cayama,
Galan sin ventura, Faisan, Cigliena cabeza pelada, Tuyuyu, Fraile (Spanish); Passarao, Cabeca seca
(Portuguese); Nengrekopoe, Amerikaanse Nimmerzat, Kaalkopooievaar (Dutch)

IDENTIFICATION
The American Wood Stork is a large (85-113 cm tall) white-plumaged, wading bird with black in the
wings and tail, a naked blackish head and neck, and a rather heavy, downcurved bill.
In its adult plumage, the head and most of the neck are featherless and covered with scaly, almost
plate-like skin, which forms a shield-shaped cap on the top of the head. The face and neck skin is
blackish grey except the cap, which has a lighter bronze tinge to it, and the nape behind the cap, which
is black. The bill is stout at its base and initially straight before tapering to a downcurved tip; it is
noticeably thicker and shorter than the bills of the Old World species of wood stork. The bill is dusky
grey with tan or reddish brown sides. Flight feathers, some coverts, and tail feathers are black with a
greenish or purplish sheen. The legs are black, but the slightly webbed toes are pink. The iris is dark
greyish brown. Males and females have similar plumages, but the male is larger (c. 3kg) than the
female (2 kg); females also have a smaller head and thinner bill.
A complete moult occurs before the nesting season. The breeding plumage features a pinkish orange
tinge on the underlining of the wings and pure white plume-like undertail feathers (coverts), which
extend beyond tail in flight. During courtship, the bill is shiny and appears polished or oiled. The feet
and lower legs turn rose; this is probably due to vascularization, as the feet pale rapidly after death
(pers. obs. M.P.K.). The legs often appear white, owing to the deposition of excreta for cooling when it
is hot at the nest (Kahl 1963b).
Nestlings are covered with a sparse grey down at hatching and weigh about 62 g. At about 10 days,
the first-down is replaced by a thick woolly white down, excepting the forehead, lores, chin, and anterior
throat, the skin of which are basically naked. By the sixth and seventh week the downy plumage turns
smoky grey on the head and neck, darker on the top of head and paler below; there is a white V-shaped
area on forehead at about time of fledging. The bill is pale yellow; the legs and feet are pale flesh-
coloured; the iris is brown. Growth and development of nestings is described more fully in Kahl (1962).
The juvenile plumage is much like the adult's but duller, and with feathering on the head and neck.
The base colour is a dull, greyish white, with some wing coverts tipped with brown. The back of the
head and upper neck are scantily feathered with coarse, smoky and buff hair-like feathers that are
darkest and longest on the top of the head. The tail and flight feathers are brownish-black, with less
sheen than in adults. As feathers are lost on the head, the skin darkens and begins to corrugate,

juveniles having less defined scaling on the naked skin. The
bill is initially pale straw-yellow and contrasts strongly with
the dark feathers of the head; the bill and legs darken
gradually with age. The bird remains in juvenile plumage
for the rest of the first year, with subadult plumage (feather-
ing on neck, bill somewhat pale) emerging in the fall moult;
some feathering remains on the neck into the third year.
Nesting first occurs in full adult plumage in their fourth year
(J.G. Ogden pers. comm. 1990).
The adult American Wood Stork is generally silent, ex-
cept during displays at the nest. In courtship, adults snap
and rattle their bills, clatter their bills together when copu-
lating, and give a rasping, hissing 'fizz' call during nest
relief. Especially in the nesting season, their wings make a
loud 'Woofing' sound during flight and landing. Large fly-
ing young give a flight call that is a trisyllabic goose-like
sound. Nestlings utter a loud Begging Call that is a high-
pitched, rasping, nasal braying 'Nyah-nyah', which gets
lower-pitched and hoarser with age. After 2 weeks, the call
becomes a loud nasal 'Wonk' (J.A. Rodgers pers. comm.
1988). When distressed (e.g. cold, fearful, very hungry)
nestlings give a short, repeated 'hiccoughing' call (pers. obs.
M.P.K.).
A large bird, the American Wood Stork flies with slow
audible wing beats; the flapping rate averages 177/min
(w = 44) (Kahl 1971b). Upon taking flight the neck is
stretched out and down and legs left to dangle, until both
can be fully extended for flight. In flapping flight, it alter-
American Wood Stork 37
nates 7-10 flaps with a short glide. It commonly uses ther-
mals to rise to great heights, from which it glides until it
picks up the next thermal. Thermal soaring and gliding are
normally used for all long-distance travel, with low-level
flapping flight used only for short journeys or when thermals
are unavailable. Because thermals are dependent on the
heating of the earth's surface by the sun, soaring/gliding
flight is restricted to the warmer times of the day, generally
between 09.00 and 16.00 h (Kahl 1964). Near the Cork-
screw colony, soaring birds have been followed aloft in a
small plane to 1525m (Browder 1984). Especially when
descending from high altitudes, they may perform remark-
able aerial acrobatics, diving steeply at high speeds and
flipping over and over (pers. obs. M.P.K.). The rush of air
through their wings sounds like a high-flying jet plane. Days
with strong, gusty winds seem to stimulate these aerial high
jinks.
Distinguishable from most other wading birds by its
height, stocky build, and downcurved bill, the American
Wood Stork is larger than ibises and smaller than the Jabiru
Stork which has all white wings and tail and massive
upturned bill. The Maguari Stork has a similar flight pat-
tern and size, but has a fully feathered white neck and head
and a less conspicuous, forked, black tail and conspicuous
white undertail-coverts. In flight, entirely black flight
feathers are diagnostic in contrast to the black-tipped pri-
maries of the smaller American White Ibis. In soaring
flight, the American Wood Stork looks very similar to the
American White Pelican (Pelecanus erythrorhynchos) but is
identifiable by its extended neck and legs. In tropical Amer-
ica, it might possibly be confused at a distance in flight with
a King Vulture (Sarcoramphuspapa); again the long legs and
neck of the stork are diagnostic.
DISTRIBUTION AND POPULATION
The nesting range includes the southern United States (pri-
marily in Georgia and Florida; isolated breeding records
have occurred further north and west (Hamel 1977, J.L.
Dusi and R.T. Dusi 1968a), southward discontinuously
through Cuba, Hispaniola, and both coasts of Mexico and
Central America (although confirmation of breeding
appears to be lacking for Guatemala and El Salvador). The
species also occurs throughout the lowlands of much of
South America. West of the Andes, it ranges to western
Ecuador and Peru. East of the Andes, it extends from the
Magdalena Valley, northern Colombia, south through
Amazonas, to eastern Peru and Tarapaca in northern Chile,
eastward through Bolivia to Cordoba, Santa Fe and north-
ern Buenos Aires in Argentina, north through Uruguay,
Brazil, and the Guianas to Venezuela (Palmer 1962:511-
512, Blake 1977:188).
American Wood Storks frequently occur outside their
usual breeding range, especially during post-breeding popu-
lation movements. The extent of these vary from year to
year. Basically, they can be divided into two classes: 'hard-
ship movements', which are most extensive after nesting
failure when ecological conditions in the breeding range
may be poor; and 'dispersal', comprising mainly juveniles
38 American Wood Stork
from that season's reproduction. The usual northern disper-
sal limits extend from North Carolina through Tennessee to
Arkansas. However, the actual limits of records reach north-
east to New York, Massachusetts, Rhode Island, Maine,
and southern New Brunswick, and north and northwest
through southern Ontario, Michigan, Illinois, South Dak-
ota, Montana, and Idaho. An extremely northern occur-
rence was recorded in northwestern British Columbia (in
1970).
Most temperate populations undergo some migration sea-
sonally. Northern-nesting birds from Georgia and northern
Florida move south into central and southern Florida in the
winter (Comer et al. 1987) and are present during the time
of the initial nesting stages of local birds. In summer, storks
that nested in southern Florida move northward to northern
Florida and the coastal plain of Georgia and Alabama.
Birds from eastern Mexico move north into Louisiana,
Arkansas, and Texas in summer; birds from western Mexico
move into southern California and southern Arizona. A
similar movement probably occurs in southern South Amer-
ica with birds moving south to Chubut, Argentina, in sum-
mer and autumn (Hudson 1920, Yamasita and dePaula
1986). In general, they can be expected to avoid situations
of extremely high water or drought, which make feeding
difficult.
The American Wood Stork is the most abundant and
widespread of the three American storks, and the popu-
lation status of this species is well-understood. During the
late 1980s, between 150-1450 pairs have nested annually in
southern Florida, and a total of 5000-6000 pairs throughout
the entire southeastern United States (Kushlan and Frohr-
ing 1986, Ogden et al. 1987, J.C. Ogden, pers. comm. 1990).
Several thousand storks are resident in Quintana Roo, Mex-
ico, and colonies of 8000-10000 pairs occur in the Usuma-
cinta Delta in Campeche, Mexico; between 1971 and 1979,
several smaller colonies totalling about a thousand pairs
were found on the west coast of Mexico (Knoder et al. 1980,
Sprunt and Knoder 1980, Luthin 1987). In Quintana Roo,
Mexico, 548 nests were counted by Cristina Ramo and
Benjamin Busto (pers. comm. 1991). A colony of 3000 pairs
was reported in Guanacaste Province, Costa Rica (M.
Gochfeld persl comm. 1989); and smaller colonies have been
found in Panama and Surinam (Spaans 1975b, Luthin
1984a). Large populations occur in the extensive freshwater
marshes of South America, being common in the Pantanal
of Brazil (Luthin 1987) and numbering nearly 6000 in the
Llanos of Venezuela (Ramo and Busto 1984). An additional
3000 were censused on the Venezuelan coast.
Large nesting colonies in southern Florida, particularly in
the Everglades and the largest at Corkscrew, in the Big
Cypress Swamp, once constituted the most abundant North
American breeding segment, numbering about 10000-
12000 pairs in the 1930s (probably not 50000-75000 pairs
as often reported) (Kushlan and Frohring 1986). There
were still over 10000 pairs in Florida in 1960 (Kushlan and
Frohring 1986, Ogden et al. 1987). However, changes in the
hydrological conditions of the wetlands have resulted in a
75% decrease in southern Florida from 1967 to 1981-82.
Population decreases have also occurred elsewhere. In
Belize, colonies known in the 1970s no longer are active,
probably owing to disturbance (Luthin 1987). The species is
considered to be threatened at the southern edge of its
range, near Sao Paulo and Rio Grande do Sul, Brazil
(Luthin 1987). Ogden et al. (1987) believe that the nesting
population in the United States shifted northward into more
but smaller colonies, expanded its range, and, perhaps,
increased in size. However, previous censusing inaccuracies
and omissions make such a shift, from south to north Flor-
ida very uncertain (J.A. Rodgers pers. comm. 1990).
The Wood Stork is one of the few storks or ibises to have
had population genetics studies completed. Stangel et al.
(1990) found that there was considerable genetic similarity
among birds nesting in different colony sites in Florida.
ECOLOGY
The habitat requirements of the American Wood Stork are
quite restricted. It is primarily a bird of freshwater wet-
lands, including inland marshes, swamps (especially
cypress swamps in the USA), lake-edge marshes and river
bottoms, but also mangrove swamps and lagoons along the
Neotropical coast. It feeds in shallow pools in inland and
coastal wetlands. The wetlands used by American Wood
Storks are characterized by seasonal fluctuation of surface
water depth. In the rainy season, these wetlands flood, but
in the dry season water levels fall, which increasingly re-
stricts water to pools and ponds where fishes become highly
concentrated.
The American Wood Stork is highly gregarious, roosting
and nesting in colonies and almost always feeding in small
groups or in mixed aggregations with other species of wad-
ing birds. Interactions among birds in an aggregation are
not common, but intraspecifically the storks use Forward
Displays in defence of a feeding site or individual distance.
At high intensities, the Wood Stork shakes its head from
side to side during the display (Kahl 1972b). Such threats
can progress to grabbing at an opponent and in bill duell-
ing. On rare occasions, birds have been seen pirating food
away from other species, such as the Great Egret (Egretta
alba) (pers. obs. M.P.K.).
The American Wood Stork feeds by tactile Groping
(Kahl and Peacock 1963, Kahl 1964) in water usually less
than 50cm deep. In trials on captives, birds temporarily
deprived of frontal vision fed as efficiently as those with
unaltered vision. It Stands or Walks slowly with its bill
placed in the water and gaped 7-8 cm. It Foot Stirs by
pumping the foot up and down on the substrate near the
Groping bill (Rand 1956) and may Wing Flash on the same
side as the pumping foot; it then repeats with the other foot
and wing, sometimes taking a step or two and turning in a
circle. While Walking, it will withdraw its bill from dense
vegetation but keep it submerged in open water. The Foot
stirring and Wing Flashing constitute a disturbance tech-
nique, and when potential prey touch the open bill, it closes
within approximately 25 ms. When a fish is caught, the
stork raises its head, bites at a large prey item, and then
swallows. While most feeding is diurnal, the birds do also
forage at night, and there is a fair bit of flight activity in and

out of breeding colonies on moonlit nights (pers. obs.
M.P.K.).
At various times many kinds of food have been reported,
including fish, prawns, crayfish, frogs, young alligators,
woodrats, young birds, crabs, snakes, turtles, and aquatic
insects and their larvae. Early reports of vegetable matter
were thought to be inadvertent inclusions, not eaten pur-
posely; however, in the 1980s a number of dead nestlings
and adults were found with impacted stomachs containing
plant material (peat) (J.A. Rodgers pers. comm. 1988).
Birds have even been reported eating cow and buffalo dung
(Kahl 1964, J.A. Rodgers pers. comm. 1990), and during
the drought of 1988 dead chicks had cow dung in their
stomachs (M. Coulter pers. comm. 1990).
Although a wide range of prey is taken, in any one
location the bulk of the diet is limited to the few types and
sizes of prey that its Groping technique can capture. In the
Everglades, we found that storks ate only fish, with 72% of
the biomass being composed of sunfish (Lepomis sp.), flagfish
(Jordanella floridae), sailfin mollies (Poecilia latipinna) and
marsh killifish (Fundulus confiuentus) (Ogden et al. 1976).
Similarly, in Georgia, 96% of the biomass of food collected
from nestlings consisted of sunfish, chubsuckers (Erymyzon
sucetta), and redfin pickerel (Esox americanus Gmelin)
(Coulter 1987b). The size of prey taken in Georgia was
consistently larger than that taken in southern Florida (M.
Coulter pers. comm. 1990). In Quintana Roo, Mexico, they
ate only fishes, primarily cichlids (Ramo and Busto pers.
comm. 1991).
In general, the American Wood Stork's peculiar feeding
technique results in its catching relatively large individuals
of whatever fishes are the easiest to catch. In southern
Florida, we were able to compare the prey taken to those
available, and we found that the storks were able to capture
selectively the species that dominated their diet, but were
relatively poor at capturing other species, including the
abundant but tiny mosquitofish (Gambusia qffinis).
The energy requirements are well known (Kahl 1962,
1964). An adult requires approximately 450 kcal/day, which
is supplied by about 520 g offish. A nestling requires about
16.5kg of food to fledge, so that an average stork family
requires 200 kg of live food or 220 000 kcal per nesting sea-
son. A colony of 6000 nests requires over one million kilo-
grams of fresh fish during the nesting season. The energy
requirements of the more typical smaller colony are con-
siderably less, of course.
To forage effectively, the American Wood Stork requires
extremely high densities offish, so in the heart of their range
they nest during the dry season when falling water levels
concentrate fish in relatively deeper pools within the marsh
or swamp (Kahl 1963a, 1964, Kushlan 1976b). In the Ever-
glades and Big Cypress areas of extreme south Florida, we
found that they selected feeding locations where fish were
being concentrated over sites with lesser densities of fish.
The connections between falling water, feeding, and nesting
are so precise that the timing of stork nesting is correlated
with the speed at which water level falls in the dry season,
and any appreciable rise of water level will result in higher
nestling mortality or cause adult storks to abandon their
American Wood Stork 39
nesting attempt (Kahl 1964, Kushlan et al. 1985). As a
result, nesting success depends on progressive drying for the
entire 4-month nesting season. Similar relationships be-
tween nesting and rates of water-level decrease have been
found elsewhere in Florida (E. S. Clark 1978a). And in that
the American Wood Stork nests at similar stages of the local
annual hydrological cycle in Central and South America
(Thomas 1985), it is likely that nesting depends on falling
water levels —and the concurrent increased food
availability—throughout its range.
To maintain suitable feeding opportunities, American
Wood Storks may use many feeding sites during the course
of the drying season, following falling water tables to a
succession of suitable foraging sites (Ogden et al. 1978,
Browder 1984, Kushlan et al. 1985). In southern Florida,
we have followed Wood Storks that were travelling by
soaring flight to feeding sites 140 km from their colony,
although 70 km is the more usual limit; in Georgia the
average distance flown was about 13km, and over 85%
were within 20km (Bryan and Coulter 1987a,b). Storks
tend to feed closer to the colony site when young are small,
returning to the nest more frequently than later in the
nesting cycle. Storks travel to nearer sites by flapping, and
more distant sites by soaring and gliding. Thus, the nesting
success of the American Wood Stork depends on finding and
obtaining the appropriate quantity and quality of food,
which in many cases is made available by fluctuating water
levels (Kahl 1964, Kushlan et al. 1975, Browder 1978,
1984).
The American Wood Stork is characteristically seen
standing motionless, alone or with other waterbirds, in trees
or on the ground and walks in an erect, 'dignified', sort of
way (thereby the name Preacher). Its overall appearance
may be summarized as stately but dull.
During periods of high ambient temperature, storks
actively excrete a dilute urine onto their legs, and the evap-
oration of this fluid acts to cool the blood in the legs, and—
through circulation—in the body (Kahl 1963b). Such be-
haviour is seen most commonly during the breeding season,
when birds are in the tops of trees for long periods. Parents
bring water to their young in the nest, enabling them to cool
themselves in the same manner. This cooling mechanism
was first demonstrated in the American Wood Stork, but it
has also been shown to occur in all other storks (as well as in
New World vultures, anhingas and some cormorants, but
not in the Shoebill).
BREEDING
The timing of the dry season differs in various parts of the
American Wood Stork's range. In southern Florida, histori-
cally, they used to begin nesting mostly at the beginning of
the dry season, in November-December, but more recently
they often begin in January-March, or even, although often
not successfully, as late as May. In 1988, nesting at Cork-
screw Swamp extended into July because of unusually dry
conditions. In most years, the rainy season begins in May or
June and any remaining nesting is aborted as the adult
storks are unable to find sufficiently concentrated food sup-
40 American Wood Stork
plies. In north Florida and Georgia, nesting occurs in the
spring and early summer (Rodgers 1990) when water levels
fall locally and the inhibiting effects of cold winter tempera-
tures have passed. In Central and South America the birds
nest during the season of decreasing water levels, beginning
in February in the Yucatan, Mexico (Ramo and Busto in
press). In Surinam, they nest from September to February
(Spaans 1975b).
They nest in wetlands, choosing isolated islands or
clumps of trees, usually over water. In southern Florida,
bald cypress (Taxodium distichwri) and red mangrove (Rhizo-
phora mangle) are the principal species used.
American Wood Storks nest in colonies, where they often
choose nesting sites on the tops of trees, as high as 25-30 m
in cypress trees and 10m in mangrove trees. In north
Florida, they nest in low vegetation, often under the canopy
(J.A. Rodgers pers. comm. 1988). Colony sites are tra-
ditionally reused year after year and are abandoned mainly
in cases of disturbance or food failure. Counts of as many as
32-45 nests have been reported from single cypress trees
(Phelps 1914, R.P. Allen pers. comm. 1959). The nest is a
well constructed platform of sticks and vines, usually with
fresh leaves and Spanish moss (Tillandsia usneoides) promi-
nently included in the lining of the cup, perhaps for insu-
lation (Rodgers et al. 1988). Nests vary from 45 to 90 cm in
diameter and 10 to 15 cm deep.
Courtship behaviour has been well studied (Kahl 1972b).
Breeding is initiated as storks form daytime 'bachelor par-
ties' at the future colony site. These clusters of unmated
birds move about from tree to tree—with loud wing-
flapping and Bill-rattling fights—before settling down on a
specific site. Such pre-pairing behaviour may last for weeks
before nesting actually starts. Eventually, display sites are
established by males and defended against both sexes by
Aerial Clattering and Forward Clattering; bouts of Display
Preening are frequent. If a mate is not obtained, males may
abandon initial display sites in favour of new trees, espe-
cially when attracted by the mating activities of other pairs.
Arching displays are given in response to disturbance, espe-
cially to potential predators below the nests.
In most courtship displays American Wood Storks droop
and spread their plume-like undertail-coverts, and bristle
the feathers on their back. The principal advertising dis-
plays are: Display Preening, characterized by the stork's
pulling the feathers of the nearly closed wing past the bill at
a relatively high rate; Swaying Twig-Grasping; a subtle
Flying Around display; and the Bill Snap. The female ap-
proaches a displaying male in a Balancing Posture, with
wings spread open and bill Gaping. The Gaping continues
after the wings have been folded in. The male often responds
to the approach with a Bill Snap. Often females are driven
off, but they generally return repeatedly until they are
accepted by the male. Following pair formation, the Balanc-
ing Posture and Gaping cease after the female is allowed to
enter the nest, and Bill Snaps are given by both sexes upon
approach of the mate. In the Wood Stork's Up-Down
display, the bird lifts its head and neck upwards, gapes the
mandibles widely, and then lowers the head and gives a
fairly loud 'fizzing' call, but few or no bill snaps. The Up-
Down may grade into Swaying Twig-Grasping, and Bill
Snaps may occur between other displays. During Copula-
tion Clattering, the male clatters his mandibles loudly and,
at the same time, beats his bill roughly back and forth
against the female's bill.
Patterns of adult attendance at the colony site depend on
the timing of thermal updrafts. Adults may stay in the
colony until mid-morning, when they use thermals to soar
and glide to the feeding grounds. They may return to the
colony site later in the afternoon, or stay at distant feeding
sites overnight before returning on the next morning's ther-
mals.
Nest-building is carried out by both sexes, and sticks are
added to the nest throughout the breeding season. Stick
gathering—which is done largely by the males during the
early stages—is a highly social affair, with dozens of birds
converging from various parts of the colony on a single tree
to gather sticks and bicker with each other. At Georgia
colonies, males did about twice as much stick gathering as
females (M. Coulter pers. comm. 1990). At Corkscrew, in
southern Florida, stick gathering activities peaked in the
early morning hours (05.30-08.30 h), before thermals devel-
oped and the major feeding flights of the day occurred (pers.
obs. M.P.K.)
Eggs are laid at 1-2 day intervals, and are incubated by
both parents from the first egg. The finely pitted shell is a
dull or cream white, usually with considerable staining, and
the eggs are ovate to elliptical ovate or elongate ovate (Bent
1926:60). Clutch size is 2-5, most typically 3, but can vary
widely from year to year at any given location. In southern
Florida, 85 clutches (pre-1940) averaged 3.48 eggs and 60
clutches (post-1970) averaged 3.33 eggs; in central and
northern Florida, 81 clutches (pre-1940) averaged 3.05 eggs
(J.C. Ogden, pers. comm. 1990). Rodgers (1990) found that
199 clutches collected in southern and central Florida aver-
aged 3.28 eggs. In Yucatan, clutch size averaged 2.16 (C.
Ramo and B. Busto pers. comm. 1991). Incubation takes
27-32 days, averaging 27.9 days in six clutches (pers. obs.
M.P.K.).
The average weight at hatching is 62 g. However, hatch-
ing is asynchronous and there is soon a substantial size
difference among siblings, the smallest and youngest of
which is the first to die if food resources prove inadequate.
After the young hatch, one adult remains in attendance
through the third or fourth week, shading the nestlings
when necessary from the heat of the sun and protecting
them from crows, which may attack the eggs or young, or
other storks, which may attempt to take over the nest.
Both parents feed the young by regurgitating onto the
nest platform, where the food is picked by the nestlings.
Small young are fed smaller quantities and more frequently
than larger young; young 5-7 days old are fed nearly 15
times per day, whereas young 3-5 weeks old are fed about
4.5 times per day, and young 6-8 weeks old were fed about
2.5 times per day (n = 231 feedings at 4 nests) (pers. obs.
M.P.K.). Parents also bring water in their gullet, which
they drool over the nestlings, some of it going into their
mouths. Nestlings Beg with a rhythmic nodding of the head,
wings partially spread.

Nestling growth is rapid (Kahl 1962), although not so
rapid as given by Ricklefs (1968), who based his calcu-
lations on too high an adult weight. By the third or fourth
week, young have grown to half of adult size and can stand
and exercise their wings (Kahl 1962). By 49-55 days (aver-
age = 51.2 days, n = 9), they can leave the nest to practice
flying. Young leave the colony at about nine weeks,
although sometimes returning to the nest site to roost and to
be fed for several more weeks. At Corkscrew in 1959, young
at least 90 days old, which had been observed throughout
the nesting season, were still returning to their former nests
to be fed by parents. Near large breeding colonies, young
tend to gather in feeding flocks that contain few or no adults;
young from smaller colonies seem to more often associate
with adults at feeding areas (pers. obs. M.P.K.).
Recent nesting success has ranged from 2.7 young per
nest in Georgia to 0.8 in northern Florida, to 0.3 in Georgia
and of course 0.0 in many instances (Coulter 1987a,b,
Rodgers et al. 1987, M. Coulter pers. comm. 1990). The
range everywhere is great and some northern Florida colon-
ies can be as productive as the Georgia ones (J.A. Rodgers
pers. comm. 1990). Recent nesting success in southern Flor-
ida has been minuscule: in the Everglades National Park,
for example, 6 years of success and 23 years of failure be-
tween 1962-1990 (J.C. Ogden pers. comm. 1990).
The greatest period of nest-mortality comes during the
egg phase of the cycle; the second greatest period of nest-
mortality is when nestlings are between 0-14 days old. In
north and central Florida colonies, between 1981 and 1985,
86% of nestling deaths were probably due to starvation,
especially of the youngest nestling in a nest (J.A. Rodgers
pers. comm. 1990). Other factors contributing to nesting
mortality include intra- and interspecific attacks and
weather disturbances. Strange, apparently unmated, Amer-
ican Wood Storks sometimes attack occupied nests, and the
ensuing fights often result in egg or nestling mortality.
The primary reason for nesting failure is rising water
levels on the feeding ground, leading to chick starvation and
colony abandonment. In the Everglades, this rise can be as
little as a few centimetres (Kushlan et al. 1975). The
phenomenon of water level related nesting failure has been
consistently found in several portions of this species' range
(Kahl 1964, E.S. Clark 1978a, Ayarzaguena Sanz et al.
1981, C. Ramo and B. Busto pers. comm. 1991).
Predation by other birds (e.g. Fish Crows, Corvus ossifra-
gus) or mammals (e.g. Raccoons, Procyon lotor) causes the
loss of some eggs or nestlings, but this is often precipitated
by other factors such as parental desertion or drying of the
substrate below the nests (Rodgers et al. 1987, Ruckdeschel
and Shoop 1987). At Georgia colonies, mammalian preda-
tion is often a major factor and reproductive success is often
determined by the outcome of a 'race' to fledge the young
before the swamp under the nests becomes dry, thus allow-
ing predators access to the trees (M. Coulter pers. comm.
1990).
Strong storms, especially when the nestlings are small,
may result in heavy or even complete losses. When accom-
panied by severe cold, storms may kill some young directly.
However, the primary effect seems to be that of dispersing
American Wood Stork 41
the food supply through the flooding of feeding sites, thereby
causing parents to desert young they cannot feed (Kahl
1964). In recent decades, low productivity in southern Flor-
ida colonies has been due largely to man-caused ecological
disturbances (see Conservation).
TAXONOMY
Formerly called 'Wood Ibis', this is a true stork and not to
be confused with the ibises of the family Threskiornithidae.
Behavioural and morphological similarities with the three
Old World species of wood stork, formerly placed in the
genus Ibis, have resulted in the combining of all the wood
storks into a single genus, Mycteria (Kahl 1972e, 1979b).
CONSERVATION
Population decreases in southern Florida have resulted in
the American Wood Stork being placed (in February 1984)
on the List of Endangered Species in the United States. The
decrease in the Everglades and Big Cypress regions is
attributable, indirectly, to water management, which has
altered the way in which the wetlands dry in the dry season
and thereby effected the timing and success of nesting
(Kushlan 1987). This has resulted in the beginning of the
nesting season shifting from November—December, to
February-April, and the storks cannot finish before the
beginning of the summer rainy season (Kushlan et al. 1975).
American Wood Storks nested successfully in the Ever-
glades only twice between 1967 and 1987.
In the Big Cypress Swamp of southern Florida, drainage
and development have reduced the foraging area available
to storks and altered the hydrology of the wetland that
remains (Browder 1978); storks nested successfully there in
only 6 years between 1963 and 1989 (J.C. Ogden pers.
comm. 1990). Similar plans to drain or to manage wetlands
for agriculture are proposed or in place for the llanos of
Venezuela, coastal marshes of Mexico, and portions of the
Brazilian Pantanal (Luthin 1987). The potential loss of an
intermittently-used feeding site at the Savannah River Nu-
clear Plant, South Carolina, led to the successful develop-
ment of artificial feeding ponds as a mitigation measure
(Coulter 1986a-c, Coulter et al. 1987b).
Elsewhere in its range, the American Wood Stork is often
threatened by human disturbance at nesting sites. Nesting
storks in Mexico and elsewhere in Latin America are hunted
by local residents for food (Luthin 1987). Such hunting
pressure is responsible for the loss of the known nesting sites
in Belize (Luthin 1987). It is especially important to protect
the nesting area in the Usumacinta Delta, Campeche, Mex-
ico, which is the most productive population of American
Wood Storks remaining in North America (Sprunt IV and
Knoder 1980).
It should be recognized, however, that the Wood Stork is
still an abundant species over much of its natural range. In
fact, the nesting population in the southeastern USA outside
of southern Florida has expanded its range and, apparently,
increased in recent years (Ogden et al. 1987). Although
population estimates are very inexact, owing to errors in-
42 American Wood Stork
herent in aerial surveying (J.A. Rodgers pers. comm. 1990),
it appears that the total US population has at least
remained stable in the last decade. Expansion into northern
Georgia and South Carolina began in 1980 and 1981, re-
spectively. Although there was, in the past, some indication
of excessive levels of pesticide contamination in the USA
(Ohlendorf et al. 1978), there is no indication of any de-
crease in clutch size (Rodgers 1990—contrary to Ogden and
Patty 1982).
The conservation strategy for the American Wood Stork
is two-fold: protection and habitat preservation and man-
agement. The protection of colony sites from disturbance,
hunting and tree cutting is essential, as is prevention of loss
and ecological changes in its wetland foraging habitat. In
southern Florida, intensive management can create con-
ditions for successful nesting (Kushlan'1987). Loss of forag-
ing habitat around the historically large Corkscrew Swamp
colony site in Florida's Big Cypress Swamp has substan-
tially reduced the area's carrying capacity for storks. It has
been discovered that American Wood Storks will feed in
artificial sites prepared for them (Coulter et al. 1987b), but
this is an expensive undertaking. In Central and South
America, conservation must centre on reducing wetland
alteration in order to protect stork populations.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 284.
 Milky Stork
Mycteria cinerea (Raffles)

Tantalus cinereus Raffles, 1822, Trans. Linn. Soc. London, 13, p. 327: Sumatra

Other names:

Malayan Wood Stork, White Malayan Tantalus, Southern Painted Stork (English); Javaanse
Nimmerzat (Dutch); Milchstorch (German); Walengkada, Walangkada (Javanese); Kuntul kelabu
(Sumatran); Bloewo, Burong upeh (Malay)

IDENTIFICATION
Slightly smaller than the Painted Stork, standing 91-95 cm tall, the Milky Stork is an all-white bird,
except for the flight feathers of the wing and tail, which are black, with a greenish gloss. Its downcurved
bill, similarly shaped to that of other wood storks, is dull yellow and sometimes tipped with white. Its
legs are dull red to flesh colour, its bare facial skin is greyish (this may be from faded skins) to dark
maroon, blotched irregularly with black. The iris is dark brown to grey-brown. In appearance, the
Milky Stork is altogether a rather dull version of its African relative, the Yellowbilled Stork.
In courtship, the white plumage brightens, becoming suffused with a very pale cream-yellow tinge
(hence 'milky' stork). The wing-coverts and back feathers have a lighter, nearly white terminal band. In
some fresh skins the wing linings are washed with salmon-pink (H.C. Robinson and F.N. Chasen
1936:205), and breeding birds show a narrow pinkish band of bare skin along the underside of the wing
(pers. obs. M.P.K.). The white undertail-coverts become elongated and fluffy. The bill turns a deep
yellow, with a greyish tan cast on the basal third. The legs become a deep magenta. The facial skin
turns dark wine-red, with a brighter red around the eye, and a pale grey or creamy smudge above each
eye; soon after courtship the facial skin fades to a paler orange-red (pers. obs. M.P.K.). (Reports of
grey, yellow, or brown soft-part coloration are probably taken from preserved skins, and these colours
may not be as in life.) The sexes are similar, males slightly larger on average and with a relatively
thinner, longer bill.
At hatching, chicks are covered with white down, By 10-14 days the contour feathers start to appear,
and by 5-6 weeks they are fully feathered. Nestlings have a dark brownish-grey bill (Hoogerwerf
1936c).
Immature birds are greyish brown above with a white lower back, rump and tail-coverts, and black
flight feathers; the wing-lining is mixed dark brown and white. The head and neck are darker brown. At
about 10 weeks loss of the small feathers on the head starts and dark naked areas are visible on the face.
After about 3 months, the bill colour changes to a warm yellow, with a greenish yellow tip. The iris
remains dark brown, and the legs are flesh-coloured (Hoogerwerf 1936c).
The sounds produced by the Milky Stork are not noticeably different from those of the other wood
storks (Kahl 1972b). Generally silent, except at breeding colonies. A falsetto 'fizzing' vocalization,
audible for about 75 m, is given during the Up-Down display, and bill-clattering and wing noise are
also heard during other social displays. Nestlings make a frog-like croaking sound when Begging
(Hoogerwerf 1936c).
In flight, the flapping-rate averages 205/min (n = 21) (Kahl 1971b). Soaring on thermals occurs
usually between 10.00 and 14.00 h (Hoogerwerf and Rengers Hora Siccama 1937). At breeding colonies
and feeding areas, flight activities are contagious, with a take-off by one bird often followed by several
other birds nearby (pers. obs. M.P.K.).

In most of its range only the white egrets and the Asian
Openbill Stork are likely to be confused with the Milky
Stork. The egrets are smaller and entirely white, and the
Asian Openbill is smaller and easily distinguished by its
peculiar greyish bill. In the northern part of its range (Kam-
puchea, Vietnam), Milky Storks may sometimes occur with
the more common and slightly larger Painted Stork, the
latter, in adult plumage, is distinguished by the black and
white breast-band and wing-coverts, pink inner secondaries,
and generally brighter soft-part coloration. Immatures of
the two species are very similar, the Painted Stork being
slightly larger and somewhat darker in overall coloration. It
is also said (B.F. King and E.G. Dickinson 1975:50-51) that
immatures can be separated by the Milky Stork having a
paler underwing-lining, which contrasts with the black
flight feathers (whereas immature Painted Storks have the
entire underwing blackish).
DISTRIBUTION AND POPULATION
The Milky Stork was formerly a widespread resident in
Southeast Asia, including Kampuchea (=Cambodia),
southern Vietnam, the lowlands of Malaysia, and several
islands in Indonesia (H.C. Robinson and F.N. Chasen 1936,
Kahl 1979b). An old specimen collected in Thailand in 1935
was recently located (Morioka and Yano 1990). Luthin
(1987) and Verheugt (1987) have summarized its recent
status. A few birds were still to be found —in mixed feeding
flocks with Painted Storks, Lesser Adjutants and herons—at
the Grand Lac, Kampuchea, in April 1968 (pers. obs.
M.P.K.). No recent records exist from Thailand. Some
Milky Stork 45
birds have apparently recolonized Vietnam since the war of
1963-75, but their breeding status is unknown.
Until 1989, the species had not been found breeding in
Malaysia since 1935 (Madoc 1936, Collar and Andrew
1988:14). However, storks in breeding plumage were seen at
Kuala Gula in July 1984, and about 20 nests were located
there in 1989; there are an estimated 100-150 Milky Storks
in this area, and the population is thought to be increasing
(K. Brouwerpers. comm. 1990).
The Milky Stork nests on several islands in Indonesia. On
the north coast of Java, the birds have bred on two island
reserves. We saw breeding birds at Pulau Dua in 1968 and
at Pulau Rambut in the 1970s (pers. obs. J.A.H.). The
species nests in eastern Sumatra at Hutan Bakau Pantai
Timor Reserve in Jambi Province. In Riau and South
Sumatra, regular breeding is suspected but not confirmed in
coastal lowlands (Silvius et al. 1986). Also in Sumatra, the
species is regularly recorded in Lampung Province from the
Way Kambas Game Preserve. On Sulawesi, Uttley (1987)
noted adults in breeding plumage and immature birds at a
mixed colony at the mouth of the Sulowath River, but no
nests were found; there is apparently a small resident popu-
lation of Milky Storks in Sulawesi (G.M.N. White and M.D.
Bruce 1986:111). In addition, it is possible that the Milky
Stork may yet be found in Kalimantan. The historic popu-
lation centre was Indonesia, but never in the period since
the Second World War has this species been numerous; and
now it is rare everywhere.
The Milky Stork is a wanderer and disperses after the
breeding season, but the details of its movements are poorly
known. It could wander from late September to April (Ver-
heugt 1987). Probably it moves to traditional feeding areas
annually after breeding, but no doubt it will shift when
conditions alter, either due to changes in rainfall patterns or
because of disturbance or harassment. It occurs seasonally
in Bali, Sumbawa, and other areas of Java. Over 1000
individuals visit the northeast coast of West Java seasonally;
it is likely that these are largely migrants from southeast
Sumatra. In Sumatra non-breeding birds are sometimes
found in the interior of the island and up to 900 m in altitude
(Marie and Voous 1988:65).
Because so much of the Milky Stork's range is inaccess-
ible, population estimates are extremely difficult to obtain.
It is possible that the entire population is as low as 5000
birds (Verheugt 1987). [The world total of '30QOO', given in
Kahl (1987a) is a misprint and should instead have read
'3000'.] We were fortunate to see this species in two island
reserves in Java in the 1960s and 1970s, and at that time it
was thought that the total breeding population was below
sustainable levels. The bird continues to be rare, although
greater searching effort has recently found isolated rem-
nants.
Several small subpopulations maintain a precious foot-
hold in Sumatra, Indonesia. In Jambi Province, 74 active
nests were found in July 1985 within the Hutan Bakau
Pantai Timor Reserve. In September 1988, nesting Milky
Storks were observed from aerial surveys at three colonies in
South Sumatra Province (Danielsen et al. in press a);
approximately 900 adults and at least 150 juveniles were
46 Milky Stork
counted in the colonies, and there were estimated to be
about 1000 Milky Stork nests (MJ. Silvius pers. comm.
1989). The largest number of Milky Storks counted in South
Sumatra was 1587 (Silvius et al. 1986), the remanant core of
this dwindling population. In Lampung Province, 116 birds
were seen at a single site in November 1985 (Milton and
Marhadi 1985).
Uttley (1986) found 33 adults and 4 immatures at a
colony site in Sulawesi, although breeding was not con-
firmed. At Pulau Rambut, West Java, there were a mini-
mum of 14 active nests in 1984, with a maximum count of 76
adults (Allport and Wilson 1986). It is estimated that be-
tween 356 and 408 birds are resident along the northern
coast of West Java (Verheugt 1987). Approximately 100
birds are resident at Kuala Gula, northern Malaysia (Dun-
can Parish pers. comm. 1989), and some 20 pairs were
found nesting there in November 1989 (K. Brouwer and
M.G. Coulter 1990).
ECOLOGY
The Milky Stork is found primarily in lowlands, and is
especially associated with coastal mangrove swamps. It also
uses various aquatic habitats, including ponds, lakes,
marshes, tidal mudflats and saline pools, and visits fish
ponds and rice fields (B.F. King and E.G. Dickinson
1975:50). In intertidal areas, the level of the tide dictates the
stork's feeding pattern. Except for the small population near
the Grand Lac, Kampuchea, most Milky Storks seem to be
associated with salt water and mangroves.
The Milky Stork often feeds in aggregations with other
wading birds, especially Lesser Adjutant Storks and Great
White Egrets (Egretta alba). It forages by Walking Slowly
and Groping with its bill slightly open, moving it from side
to side, and also by Walking and Probing in the mud. It has
been reported to Foot Stir, like the other wood storks (Hoo-
gerwerf 1936c). The main method of feeding is by tactile
location, but undoubtedly it visually locates mud-skippers
which are abundant on the mudflats of Indonesia and Viet-
nam (Silvius 1988). On intertidal mudflats in western
Malaysia, a bird was observed to Probe in the mud, to
Grope in the water and to search visually for prey (Swennen
and Marteijn 1987). We have seen this species on Pulau
Rambut feeding in the shallow pools beneath the mangrove
bushes in the company of egrets. Here food supplies were
abundant, allowing this stork to consume sufficient food in a
short period and enabling it to lounge around for prolonged
periods of the day. At the Pulau Dua, Java, breeding colony
in March 1968 there was considerable in-and-out flight
activity on nights with a full moon, indicating that the birds
not only forage, but also visit their nests, at night (pers. obs.
M.P.K.).
The main diet is small fish, primarily mud-skippers
(Gobiidae) in Malaysia (Swennen and Marteijn 1987), but
snakes and frogs have also been recorded (Hoogerwerf and
Rengers Hora Siccama 1937). Hoogerwerf (1936c) observed
parents feeding 20 cm fish, eels and mud-skippers to their
young in the nest in western Java.
Like other wood storks, the Milky Stork will spend time
soaring on thermals, and it uses this method of flight when
moving from area to area. At high tide, birds are often seen
resting in mangrove forests. They are said to be generally
shy and vulnerable to disturbance (Verheugt 1987),
although we found that birds at the nest could be
approached easily.
BREEDING
In Indonesia, breeding takes place after the rains, during
the dry season that usually lasts until October. The species
probably depends on particular water levels for feeding; the
start of the breeding season probably coincides with maxi-
mum fish stocks, following reproduction of fish during the
rainy season. In Java, nests were often in black mangroves
(Avicennia marina) 8-30 m high (Hoogerwerf 1936c, Ver-
heugt 1987), and in Sumatra nests were located in red
mangroves (Rhizophora apiculata) (Silvius 1988). On the
north coast of Java, nests on Pulau Rambut were built on
top of the highest trees, some 30 m above the ground and we
found some of the nests in similar position on Pulau Dua
when the species still nested there. A few large trees on one
side of the island were used in preference to the lower
mangroves and creeper-covered bushes occupied by the
bulk of the other wading birds, although some Milky Storks
nested as low as 4 m at Pulau Dua. Unfortunately, since the
mid-1970s Pulau Dua has become joined to the mainland by
accumulated silt, allowing access to humans and other pred-
ators, and since that time the storks have not nested at this
site. Several South Sumatran colonies were built 2-15 m up
in mangroves within 1—4 km of the coast; here the Milky
Storks nested with Lesser Adjutant Storks, Oriental White
Ibis, and several species of herons (Danielsen et al. in press
a). Malaysian nests were 8-1 Om above the ground, mostly
in dead trees (S.H. Yatim pers. comm. to K. Brouwer,
1990).
The nests are constructed of sticks and lined with fresh
vegetation. Hoogerwerf (1936a) described the nests as simi-
lar to those of the Grey Heron (Ardea cinerea) but slightly
more robust and containing thicker branches.
We have found the breeding and courtship of the Milky
Stork to be similar to other members of the wood stork
group, although certain threat displays, such as Aerial Clat-
tering and the Forward display, have not yet been observed
(Kahl 1972b). During many displays at the nest, performing
birds retract the skin of their head and upper neck, exposing
two or three times as much area of naked skin; the ampli-
tude of this retraction is even greater than that seen in the
Yellowbilled Stork. Between displays, when the retraction is
relaxed, and the skin hangs in loose folds and wrinkles, like
wattles (pers. obs. M.P.K.).
The male advertises using Display Preening and the
arriving female reponds with the Balancing Posture and
Gaping, which may continue for several days during pair-
formation. The male uses a Flying Around display, just after
the female occupies the nest site, and Up-Down displays are
given in greeting. In the Milky Stork, the Up-Down in-
cludes four or five bill snaps in a series between hissing-
screaming vocalizations. Swaying Twig-Grasping is also a

typical display. In response to disturbances at the nest, the
Arching display is given; the photograph in Hoogerwerf
(1936c) shows it to be essentially identical to the homolo-
gous display in the other wood storks.
Allopreening between pairs is frequently followed by
body shaking. Head rubbing is another common comfort
movement; the bare skin of the head may first be oiled by
rubbing it on the preen gland and is then rubbed on the
plumes of the back. This movement is carried out quite
frequently by birds perched nearby an incubating partner.
Nesting is colonial, and, in Java, Hoogerwerf (1936c)
reported 5-8 nests in a single tree. Most colonies are rather
small, with from 10-20 up to a few hundred nests.
The eggs are chalky white and usually very dirty. At West
Java colonies, 9 clutches contained 3 eggs each (Helle-
brekers and Hoogerwerf 1967); sometimes the clutch is 4
(Hoogerwerf 1949). The incubation period is estimated at
27-30 days (Hoogerwerf 1936c).
Parents feed by regurgitating food onto the floor of the
nest, where the nestlings pick it up. Small young are fed
more often than large young; up to 4 weeks, chicks may be
fed twice per hour, whereas older chicks may only be fed
once in the afternoon (Hoogerwerf 1936c). During the heat
of the day, the young are often shielded by the open wings of
the adult which stands over the nest. Adults sometimes
bring water to cool nestlings and for them to drink (drooled
from the parent's bill). Heat stress is also alleviated by birds
excreting a dilute urine onto their legs, which turns them
white. At Pulau Rambut, Java, in 1984, active nests held
about 2 young each (Verheugt 1987). By 6-7 weeks the
young are able to leave the nest and begin to fly, and by 8
weeks they are flying well but are still fed in the nest by the
parents (Hoogerwerf 1936c).
TAXONOMY
The species was formerly and inappropriately called 'Wood
Ibis', which led to unnecessary confusion with the true
ibises (family Threskiornithidae). Because the morphologi-
cal and behavioural similarities are so great, this species,
formerly in the genus Ibis along the the other Old World
wood storks, is now included with them and the American
Wood Stork in the genus Mycteria.
CONSERVATION
This species was essentially eliminated from Vietnam by the
widespread destruction of mangrove swamps during the
Milky Stork 47
Southeast Asian War (1963-75). Extensive mangrove refor-
estation and protection by wardens may have resulted in
some recolonization in recent years. Elsewhere it is seriously
threatened by poaching, disturbances at nesting colonies,
mangrove deforestation, tidal rice cultivation, fish-farming,
and—in Indonesia—human resettlement (Luthin 1987,
Verheugt 1987, Collar and Andrew 1988:14). Of the 26 sites
at which the species is known to occur, 16 (62%) are
reported to be under moderate to severe threat (D.A. Scott
and C.M. Poole 1989). In addition, specimens appear from
time to time in the zoo trade (Danielsen et al. in press b);
some of these have been purchased as 'juvenile Painted
Storks' and, on maturity, proved to be Milky Storks instead.
At present, some 70 Milky Storks are kept by nine zoos
outside of Indonesia (K. Brouwer pers. comm. 1990).
Indonesia Interwader (the South East Asia Pacific Shore-
birds Study Programme, with headquarters in Kuala Lum-
pur, Malaysia), the ICBP and other European-based con-
servation organizations participated in wide-ranging
surveys in the mid 1980s, providing much of the status
information we report here. A Milky Stork conservation
programme has been proposed. It is especially critical to
establish an integrated network of wetland preserves, as is
being done in Vietnam. However, the loss of coastal wetland
habitat continues, and in Indonesia is estimated to account
for the loss of 100000 ha per year (Verheugt 1987). The
near total destruction of the mangrove forest in Java is the
most serious threat to the species there. Also, in many areas
birds are netted throughout the year and eggs are taken
during nesting.
The Milky Stork is now an officially protected bird in
Malaysia and Indonesia (Verheugt 1987), and as the core of
the population is confined to areas near gazetted nature and
wildlife reserves, it is hoped that the considerable devas-
tation brought about in recent years by humans can be
mitigated. Certainly without active conservation efforts,
especially those sponsored by Interwader and supported by
ICBP and other conservation organizations, this stork has
little chance of survival. The species has been listed on
Appendix I of CITES since 1987.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 285.
 Yellowbilled
Stork
Mycteria ibis (Linnaeus)

Tantalus ibis Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 241: Egypt

Other names:
Wood Ibis, African Wood Stork (English); Tantale ibis, L'ibis tantale (French); Afrikanischer
Nimmersatt (German); Kamindo (Mandingo); Tsagtsagi, Tsagagi (Hausa); Okogliech (Luo);
Gangala (Mozambique); Levlosyane (Sotho); Nepando (Kwangali); Geelbekooievaar (Afrikaans);
Naedjah, Badja (Arabic)

IDENTIFICATION
This medium-sized stork, standing 90-105 cm, appears all white, except for its black wings and tail.
The back of its head and neck are a dirty white, whereas its back, wing coverts, breast, and belly are
white, more-or-less suffused with pink on the wings. The tail, flight feathers, and greater primary
coverts are black. The distinctive yellow bill is long and slightly decurved, as is the case for all wood
storks. The head is feathered to the hindcrown, and the bare skin of the forehead, face and throat is
orange to red; the legs are pale red.
Courtship and breeding colours are much brighter. The bill becomes a rich golden-yellow, while the
face turns a bright carmine-red, grading to orange posteriorly near the feathers. During courtship
displays, the skin of the head is retracted, considerably enlarging the visible area of bare red skin (Kahl
1972b). The legs and feet become a brighter red. Much of the white plumage becomes suffused with
pink and the wing-coverts are crimson, especially the greater underprimary- and underalula-coverts,
each feather being tipped in white. When freshly moulted, the black feathers of the wings and tail are
glossed with green and purple. The sexes are alike, but the male is slightly larger.
At hatching, the nestling weighs about 60 g and is covered with a sparse grey down. By 10 days, the
weight increases to about 500 g, and the grey first-down is superseded by a white, woolly second-down.
The bill and bare facial skin start out mostly yellow and gradually change to black with age. By about
5 weeks of age the bill starts, from the tip, lightening to a pale yellow. Nestlings have dark brown irises
and legs and feet that progress from a fleshy-pink to a cyano tic-blue (pers. obs. M.P.K.).
Immature birds have the feathering on the head extending further forward than in adults. They are
greyish brown, paler underneath, with a dull partially bare face of orange and a horn or dull yellowish
bill. The legs and feet are a brown horn colour, and the primary, secondary and tail feathers are
blackish brown. By about the age of fledging, they start to develop some salmon-pink coloration in the
underwing linings (pers. obs. M.P.K.). After the first year, the body plumage is white washed with grey,
the flight feathers of the wing and tail become black, and adult-coloured soft parts begin to appear,
although the pink flush does not occur until later. Birds are probably not fully adult for 3 years.
The Yellowbilled Stork is generally a silent bird, but during social displays a rather weak hissing
scream is uttered. At breeding colonies a variety of bill-clattering and 'woofing' wing sounds are heard.
Nestlings give a loud, repeated, and monotonous braying call when Begging adults for food.
The Yellowbilled Stork flies with its neck and legs fully extended, using either flapping or soaring
50 Yellowbilled Stork
flight. The flapping rate averages 186/min (n = 20) (Kahl
1971b). For flights to distant feeding grounds, Yellowbilled
Storks soar on thermals and glide, as do most storks, to
cover long distances without wasting much energy. When
descending from on high, either at the feeding areas or the
breeding colony, birds often dive at high speeds, side-slip
and flip; they appear to enjoy this aerial play.
The Yellowbilled Stork can be confused with wintering
White Storks. However, the decurved yellow bill, black tail
and rosy pink cast of the adults are distinctive. In flight, at a
distance, it might be mistaken for a Great White Pelican
(Pelecanus onocrotalus) or an Egyptian Vulture (Neophron perc-
nopterus), both of which have shorter necks and legs and
white tails.
DISTRIBUTION AND POPULATION
The Yellowbilled Stork is widely distributed in suitable
habitat in tropical Africa, south of the Sahara from Senegal
and Somalia south to South Africa, where it is rare south of
the Orange River, and also to Madagascar. It seems, gener-
ally, to avoid the heavily forested regions of central Africa.
It is a straggler to southern Tunisia, Morocco, and Israel,
and formerly to Egypt (F.J. Jackson 1938:80, Castan and
Olier 1959, Paz 1987:34).
The species is particularly widespread in East Africa,
where it breeds in Kenya (3 sites), Tanzania (5 sites), and
Uganda (common, but not recorded breeding) (Kasoma
and Pomeroy 1987). In the Sudan it breeds at Malakol and
probably other sites. In West Africa, it nests from Gambia
to northern Nigeria, often inside walled cities (Bannerman
1953:180). It was found breeding in Zululand, South Africa,
in 1949 and again in 1954, 1960 and 1961 (Clancey
1964a:47^8, Berruti 1980b). It has bred in northern Bots-
wana (W. Fraser 1971) but is generally uncommon south of
the well-watered parts of Zimbabwe and northern Bots-
wana. It occurs sparingly in the non-breeding season over
much of South Africa, although Clancey (1964a:47) found it
present year-round in the coastal waters of Natal. It is found
quite widely in western Madagascar (Rand 1936:340); Lan-
grand (1990) reported that there is no direct evidence of
nesting there now, although in October near Lake Kinkony,
young birds hardly able to fly were seen.
The Yellowbilled Stork undertakes local movements in
Kenya and migrates from north to south in Sudan with the
rainy seasons, leaving the Sudd during the flooding period
(Luthin 1984a,b). In the Bangweulu area of northeastern
Zambia it leaves when the water is high—and fish are hard
to catch —and returns in large numbers when the swamps
begin to dry (Brelsford 1946). Although it migrates regu-
larly to and from South Africa, its general movements are
poorly known, and in some areas it remains year-round. It is
probably best to describe the Yellowbilled Stork as a facul-
tatively nomadic species, avoiding areas where water or
rainfall conditions are inappropriate.
The overall population may be considered abundant and
stable. The species is not uncommon as far north as the
Sudan, and in Kenya we found it to be the commonest
species in a colony on the Tana river, numbering about 2000
birds. In the following year after the long rains it also nested
there, some 200 birds being present. Larger numbers than
this are found in eastern and central Africa in seasons when
rainfall is adequate. A colony of at least 2000 pairs was
found in western Tanzania in February 1962 (Stronach
1968).
ECOLOGY
This stork appears to use a somewhat wider range of habi-
tats and have a broader diet than other wood storks. Its
habitat use is particularly diverse, including large marshes,
swamps, margins of lakes or rivers, rice fields, and even
small pools. It feeds in tidal pools along the beach in East
Africa (Brown et al. 1982:174). Most of its food is taken in
water about 10-40 cm deep, using its partially open bill to
Sweep and Grope.
The Yellowbilled Stork feeds by tactile location, Stand-
ing, Walking Slowly, Groping, Foot Stirring, and Wing
Flashing. It will often stand still to await the arrival of prey,
snapping its partially open bill at its touch. Foot Stirring is
often accompanied by the flashing open of the wing on the
same side (Talent 1940), as with the Painted Stork and the
American Wood Stork. Prey is quickly swallowed with a
toss of the head.
The species usually feeds singly or in small groups, but at
times in the hundreds (F.J.Jackson 1938). Birds in foraging
groups often threaten each other with Forward Displays, in
which the neck is retracted, the black tail is cocked upward,
the wings are slightly drooped, and the back and neck
feathers are erected (J. Cooper 1969, Kahl 1972b). It will
sometimes feed in company with other species, often herons,
spoonbills, or Great White Pelicans (Pelecanus onocrotalus).
On rare occasions, Yellowbilled Storks have been seen feed-
ing in the wake of a moving crocodile or hippopotamus,
apparently taking advantage of organisms stirred up by the

larger animal (Pooley 1967a). The storks feed apparently for
only a short time, obtaining their requirements often very
quickly and then resting.
Frogs, small fish, aquatic insects, worms and crustaceans,
and possibly some small mammals and birds are eaten. A
tilapia of around 150 g appears to be about the maximum
size offish that can be swallowed (Bell-Cross 1974). Sca-
venging of food regurgitated by cormorants has been ob-
served (Kasoma and Pomeroy 1987).
The Yellowbilled Stork will roost with other species on
the ground, and at night in large tree roosts. When resting
on the ground, birds often sit on their tarsi. Sometimes,
especially after a rain or following bathing, they stand with
their wings fully extended to the side until dry (Kahl
1971g).
BREEDING
The breeding pattern of this stork varies throughout the
African continent. Nesting areas are subjected to annual or
semi-annual rainfall, and the Yellowbilled Stork will nest
either at the peak of or at the end of the long rains, whenever
food is most abundant because of local ecological con-
ditions. Especially in extensive marshlands with little topo-
graphical relief, nesting is at the end of the rains, as declin-
ing water levels concentrate fish, similar to the situation
with the American Wood Stork in southern Florida. How-
ever in other habitats, such as in the marshes near Lake
Victoria in western Kenya, breeding occurs at the peak of
the rains, as fish move out of the lake and adjacent rivers
and into the flooded marshes (Kahl 1968). There, during
the dry season, the fish are in deeper waters and unavailable
to the storks; when fish move into shallower marshes to
spawn, they and their fry are most available to all fish-
eating birds, who time their breeding cycle to this annual
event. Cases of off-season breeding of Yellowbilled Storks
have been reported from western Kenya (J. Parsons 1977),
northern Botswana (Tree 1978) and eastern Kenya (Han-
cock 1984:41), when unseasonal rainfall caused local flood-
ing and ideal feeding conditions. Breeding sites and feeding
grounds are often far apart, and birds will travel quite long
distances—usually by using thermals for soaring and glid-
ing flight—to obtain food.
Nesting is colonial, often in company with other water-
bird species, but sometimes the Yellowbilled Stork is the
sole or major occupant of a nesting site. Nests are built
throughout a mixed colony, with perhaps up to 20 storks
nesting close together in any one part of the colony. The
nests may be built in small trees, usually over water, or in
larger trees over dry land. As many as 50 nests have been
counted in a single large tree (Bannerman 1953:180).
Pair-formation begins when the male displays from its
chosen position, using Display Preening which appears to
expose the brightest parts of the wing to an approaching
female. The interested female adopts a Balancing Posture,
with her head low and forward, bill agape and wings spread.
She will also use Gaping without the spread wings, a behav-
iour pattern unique to the wood storks. Aerial Clattering
and Swaying Twig-Grasping are characteristic. In many
Yellowbilled Stork 51
displays, the skin of the head is pulled back, allowing the
bright-coloured bare skin to expand over the entire head.
Up-Down displays are performed by the pair. Bill clattering
occurs during copulation in a similar manner to other wood
storks, and threat displays and anxiety movements are simi-
lar to those of the other members of this genus.
In traditional sites, old nests—if any remain from a pre-
vious season—may be repaired, but usually new stick nests
are constructed and lined with grass or green leaves. Nests
take up to 10 days to build and are 80-100 cm in diameter
and 20-30 cm thick. 2-4 eggs (usually 3) are laid; average
clutch size (n = 45) was 2.5 (Brown et al. 1982:175). They
are a dull white to greenish white, usually unmarked but
rapidly become stained. J. Parsons (1977) found that there
was a significant decrease in 'the width of Yellowbilled
Storks' eggs from a Kenyan colony as the season progressed.
Both parents share duties of incubation, brooding, guard-
ing and feeding the young. Incubation takes about 30 days.
Young hatch at 1- or 2-day intervals and grow rapidly.
Their insatiable appetites make the German common name
'Nimmersatt' (never satisfied) particularly appropriate.
One parent usually attends the young constantly until they
are about 21 days of age, at which time both adults forage
much of the time to fulfil the nestlings' food demands. At
this age, young suddenly become quite agressive, and this
may be in response to the need to defend themselves from
attack, particularly from other Yellowbilled Storks that
attempt to take over the nest when adults are absent.
Feeding is by regurgitation on to the nest floor, and water
is passed from adults (Stronach 1968) by dribbling it into
the nestling's bill. Adults will shade the young with open
wings, but brooding of young quickly ceases. The slower
movements of these storks when nestlings reduces losses by
falling from the tree, unlike heron species where fighting and
rapid early movements away from the nest result in heavy
losses.
Generally, breeding success seems to be relatively high,
with broods of 1-3/nest recorded in East Africa (Brown et
al. 1982:175). Egg predation by Fish Eagles (Haliaeetus
vocifer) was heavy at a colony near Kisumu, Kenya, in 1975.
At this colony, at least 236 eggs were laid in 95 nests
(average of 2.48 eggs/nest). An estimated 61% hatched and
38% were predated; only 31 young fledged, giving a success
rate of only 0.33 young/nest (J. Parsons 1977).
The young take their first flights at 50-55 days and soon
leave the nest and become fully independent. Juveniles take
at least 3 years to attain adult appearance and may not
breed until later; firm data are lacking however.
TAXONOMY
In the past, the Yellowbilled Stork, along with the Painted
and Milky storks, was classified in the genus Ibis. However,
intensive studies have shown that they are closely similar to
the American Wood Stork, and so all four species have been
united in the genus Mycteria (Kahl 1971f, 1972e).
In the older literature, storks of the genus Mycteria (in-
cluding the former genus Ibis] were often called by the
common name of'Wood Ibis'. This led to much confusion
52 Yellowbilled Stork
with the true ibises (family Threskiornithidae). It has long
been recognized that these birds are, indeed, true storks
(Garrod 1875a,b) and should not be called ibises. Names in
current use reflect that fact and avoid much of the confusion
of the past. However, readers consulting the older literature
should be aware of these nomenclatural changes.
CONSERVATION
This is a widespread species, which appears capable of
responding to short-term natural changes in habitat con-
ditions. In East Africa, it is both abundant and stable but is
subject to pressure from poaching and from reduction of
habitat. Whilst increasing human populations in many
parts of Africa constitute a long-term threat, it cannot be
considered other than an abundant species with a compara-
tively large, stable population at present.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 286.
 Painted Stork
Mycteria leucocephala (Pennant)

Tantalus leucocephalus Pennant, 1769, Indian ZooL, p. 11, pi. 10: Ceylon (Sri Lanka)

Other names:

Rosy Wood Ibis, Indian Wood Stork, Pelican-ibis (English); Indischen Nimmersatt, Buntstorch
(German); Tantale indien (French); Janghil, Dhok (Hindi); Kankari (Mirshikars, Bihar); Kat-
sarunga, Ram-jhankar, Sona-jangha (Bengali); Lamjang, Lungduk (Sind); Chitroda (Kutch); Yeru
kald-konga, Yerri kali-konga (Telugu); Ghenga narai, Singa nareh (Tamil); Sangu-valai-narai,
Changa vella nary (Tamil, Sri Lanka); Dae-tuduwa (Sinhalese); Hnet-kya (Burmese); Lai-tou-kuan,
Tu-Tsae-kuan (Chinese)

IDENTIFICATION
The Painted Stork is a medium-sized, conspicuously patterned, black and white stork, standing 93-
100 cm tall. The head is partially naked and orange; the bill is yellow and slightly decurved at the tip.
The face and crown of the adult is featherless, and the skin is an orange-yellow to reddish orange
colour. The nape, neck, back and breast are white, with a black-and-white band horizontally across the
breast. The primaries, secondaries and tail are black. All the wing-coverts (except the upper greater
secondary coverts, which are white) are black, glossed greenish, but feature prominent white terminal
bands. These 'chequered' areas of the wings and breast are one of the distinguishing features of the
species. The innermost secondaries and their coverts, close to the tail, are rose pink. The legs and feet
are also pink to dull red, but they often look white from the excreta splashed over them, especially
during the nesting season. The eyes are dark brown. The sexes are alike, but the male is slightly larger.
In the breeding season, the plumage becomes neater and cleaner looking, with the black-and-white
coverts and breast-band contrasting strongly with the surrounding feathers. The elongated inner
secondaries and their coverts become a brighter pink, with white terminal bands. The bill colour
deepens, and the face becomes reddish orange with a glaucous cast and the throat pinkish. By retraction
of the skin, the extent of the bare head increases during some displays, but not as dramatically as in the
Yellowbilled Stork of Africa. The legs turn a brighter reddish magenta, and the iris becomes a paler
grey-brown (Kahl 1972b, Breeden and Breeden 1982).
At hatching, the nestling is partially feathered with a pale greyish down. Partially grown chicks are
covered in white down, accentuating their naked, jet black faces. The black bill is tipped with pale
yellow, more on the upper than lower mandible (S. Ali and S.D. Ripley 1968:95). Immature birds have
greyer plumage, with the head and neck pale brownish, arid the back darker with lighter buffy greater
secondary coverts. They lack the band across the chest and the pink secondaries of adults. Head
feathering of the young is lost gradually. The forehead and anterior of the crown become featherless by
one year, and the entire head by two years of age. This feather loss is accompanied by an increase in
lipoid secretion, probably to protect the exposed skin (Shah et al. 1977).
Adults are generally silent away from the nest. During displays, the typical wood stork 'fizzing' call is
given, but it is somewhat weaker in this species (Kahl 1972b). Also wing-woofing and various forms of
bill-clattering are heard during displays, fights and copulation. Nestlings vocalize loudly when Begging
to be fed; their voices are not noticeably different from the other wood storks. Year old juveniles can still
vocalize loudly, but become practically voiceless, as adults, by 18 months (Shah and Desai 1972).
In flight, the Painted Stork flies with neck extended and slightly downcurved. The flapping rate

averages 190/min (n = 70) (Kahl 1971b). The Painted Stork
readily soars high on thermals, and can be seen travelling
long distances with other soaring species.
In most of its range only the white egrets and the Asian
Openbill Stork are likely to be confused with the Painted
Stork. The egrets are smaller and all white, and the Open-
bill is smaller and is easily distinguished by its peculiar
greyish bill. In parts of southeast Asia (e.g. Kampuchea,
Vietnam), it may occur alongside the slightly smaller Milky
Stork. Adults are easily distinguished by the more intricate
black-and-white pattern on the wings and across the breast
of the Painted Stork. It is also said (B.F. King and E.G.
Dickinson 1975:50-51) that immatures can be separated by
the Painted having the entire underwing blackish, whereas
immature Milkys have a paler underwing-lining, which
contrasts with the black flight feathers.
DISTRIBUTION AND POPULATION
A lowland species entirely dependent on water, the Painted
Stork's range extends throughout the Indian subcontinent
from Pakistan to Sri Lanka and eastward as far as south and
eastern China, Thailand, Kampuchea and Vietnam.
Although widespread in India, it is more localized elsewhere
in its range. A small nesting colony remains in the Indus
Delta of Pakistan, and a few birds appear erratically as far
as Mainwali in the Punjab (T. Roberts pers. comm.). Nu-
merous Indian colony sites are known from Gujarat, Saur-
ashtra, Rajasthan, and elsewhere (Luthin 1984a). Colonies
are located within the Colombo Zoo in Sri Lanka and the
Delhi Zoo in India. There are no recent breeding records in
Bangladesh (M.A.R. Khan 1987). In 1982-83, Painted
Storks were observed in Burma. In Thailand, there is an old
nesting record from Ban Yang (J.H. Riley 1938), and the
only known recent nesting site is at the Thale Noi Non-
Hunting Area, in southern peninsular Thailand, where only
one pair remained in 1988 (P. Round et al. unpub. report).
Four small colonies are known from Vietnam, in the coastal
wetlands. Non-breeders occasionally wander southward in
Painted Stork 55
the Malay Peninsula, as far as Kuala Lumpur (Gibson-Hill
1949:29).
The species is non-migratory, moving nomadically after
nesting to wet areas, dispersing in small parties to wherever
food is available.
The Painted Stork remains locally abundant in India and
Sri Lanka, but elsewhere its numbers have been drastically
reduced. One of the largest known colonies—the Keoladeo
Ghana, near Bharatpur, Rajasthan, India—contains several
thousand pairs in a good year (S. AH 1953:535, Kahl 1974).
A colony in Baunagar, Gujarat, India, numbered 170 nests
in 1983 (Luthin 1984a). A census in midwinter 1989, re-
vealed a total of nearly 5000 birds in India and 850 in Sri
Lanka (D.A. Scott and P.M. Rose 1989). The Pakistani
population is small, and the species is now very rare in
Bangladesh (M.A.R. Khan 1987), Burma, Thailand and
Vietnam (Luthin 1987). It is unlikely that any substantial
breeding occurs in any of these areas today. Earlier this
century, it was said to be abundant in northwestern Kam-
puchea (Cambodia) (Delacour 1929); a few birds were seen
there in April 1968, including some half-grown young, being
held captive for human-consumption (pers. obs. M.P.K.),
but nothing is known of its status now. In southern Viet-
nam, where it was common before the war, it now breeds
only in four small colonies of 5-10 pairs each (Luthin
1984a); its present status there is uncertain. In China, it
formerly occurred in summer in the southeast (La Touche
1931-34:441; Etchecopar and Hue 1978:71); it is almost
certainly extirpated there now.
ECOLOGY
Painted Storks are found in freshwater marshes, ponds,
flooded fields, sometimes river banks, and rarely on the
coast and in salt pans. In the non-breeding season, they
have been observed feeding in relatively dry agricultural
fields (AJ. Urn pers. comm. 1990).
It is a tactile feeder, searching shallow waters with its bill
half open, Groping and making use of Foot Stirring and
Wing Flashing to disturb its prey. Birds Walk Slowly, Grop-
ing as they move, and when food is plentiful they will feed in
quite large parties. During Foot Stirring, the wing is often
flicked quickly open on the same side in order to direct fish
back toward the open bill. Only occasionally do feeding
birds perform Forward Displays at neighbouring birds. The
bilLis held forward, with the body near horizontal and the
neck retracted; the feathers on the back, neck and breast are
erected, undertail-coverts drooped and tail cocked. If the
opponent does not retreat, its bill may be grabbed and brief
grappling takes place.
Their diet is almost entirely fish, although S. AH and S.D.
Ripley (1968:93) reported that Painted Storks will also eat
reptiles, frogs, crustaceans and insects. The length offish
consumed by adults is usually 5-25 cm. In Delhi Desai et al.
(1974) found that the fish taken were mainly carp and
catfish during the storks' nesting season, which is also the
time at which these fish breed and are most available.
Stomach contents also showed that besides fish, these storks
had consumed fragments of plant fibre, vegetable debris,
56 Painted Stork
small stones, aquatic insects and one small frog. It is esti-
mated that an adult bird consumes as much as 660 kcal, or
600 g of fish, per bird per day to maintain body weight
(Desai et al. 1974); a family of storks consumes approxi-
mately 240 kg of fish during a breeding season. In Delhi,
breeding is in the last phase of the rainly season, when fish
are available in large quantities (Desai et al. 1977).
After feeding, Painted Storks frequently stand on the
bank in a hunched posture (S. Ali and S.D. Ripley 1968:93).
Standing birds often rest on one leg and clasp the tarsus of
that leg with the other foot (Henry 1971:384). When resting
on the ground away from the nest, they sit on their heels
with their tarsi folded forward horizontally.
BREEDING
A colonial breeder, the Painted Stork often nests in mixed
colonies of storks, ibises, spoonbills, cormorants and herons.
In the larger colonies in India, such as at Bharatpur, up to
several thousand pairs nest. Colony sites are traditional and
are often used for many years if undisturbed (S. Ali and S.D.
Ripley 1968:93). Breeding normally starts during the latter
part of the rainy season, but its timing is dependent on
water levels and food; if the rains fail, breeding will not start
or will be abandoned for the year.
Old nests may be repaired, if available, or new platforms
are built (Desai et al. 1977), in trees or tall bushes, often
standing in or overhanging water. Some colonies consist of
nests of only Painted Storks, in a single large tree such as a
peepul (Ficus religiosa); we know of several such colonies, in
northern India, in the centre of villages. At the large colony
near Bharatpur, India, nests are in many low, scattered
babul trees (Acacia arabica). Nests have also been placed in
trees such as kandi (Prosopis spicigera), keli-kadamb (Stephe-
gyne parviflora), and Barringtonia racemosa (S. Ali and S.D.
Ripley 1968:94). For 15 nests in Delhi, it took 4-8 days
(average = 6.3 days) to build the nest. Sticks 30-60 cm long
are used for construction, with green leafy material as the
lining. Males do most of the stick gathering, making 5-6
trips per hour in the morning and afternoon (Desai et al.
1977). The nest itself is a rather flimsy stick platform some
55-80 cm in diameter, lined with a small amount of leaves,
straw and water weed. Fresh vegetation or leafy twigs are
added throughout the occupation of the nest.
Groups of unmated males often settle only temporarily on
nest sites, and if they do not acquire mates, they move on to
another tree taking unmated females with them. Such
'bachelor parties' (Kahl 1972b) may contain a dozen or
more males and an equal number of females, and they form
a conspicuous feature of any colony of wood storks at the
beginning of the breeding season.
Painted Storks, like other wood storks, perform a number
of ritualized displays at or near the nest (Kahl 197la). In
order to form pair bonds, the male stork occupies a potential
nest site while Display Preening, and is approached by an
unmated female. Displays are often of a high intensity and
are more frequent at the commencement of breeding. In
Display Preening the stork pretends to strip down its wing
feathers with its bill, in a rhythmic pattern. The female
reponds by landing near, assuming the Balancing Posture,
with wings widely spread and bill open, in a display called
Gaping. She may maintain this posture for a half of a
minute or more, before closing her wings, although still
Gaping.
Males defend territories with Aerial Clattering, Forward
Clattering and Bill Snaps. Such rituals are common to all
the wood storks, as is the Arching display given usually at
the approach of a possible predator below the nest site, and
which is often copied by nearby birds and spreads across the
colony. In the Bill Snap, given by both sexes during pair
formation, the bill is lifted to the horizontal and a single
audible snap is given. Once a female has been accepted at
the nest, the male performs a Flying Around display, circ-
ling around and landing again next to the mate. Up-Down
displays usually follow such flights, as they do whenever a
member of a pair returns to the nest site. The birds swing
their heads and necks upwards to near the vertical, with the
bill open. They then lower the head, swinging it from side to
side. The Up-Down of the Painted Stork is similar to the
other wood storks except that its softer hissing vocalizations
are interrupted by double or triple Bill Snaps as the head is
lowered. After pair-formation is complete, returning birds
carry out mutual Up-Down displays together at the nest.
In the Painted Stork, the Swaying Twig-Grasping plays a
particularly significant part in early courtship, being espe-
cially ritualized in this species. On the nest-platform the
bird bends over and touches or grasps lightly at twigs at
intervals of 1-2 s. Sometimes the neck is swung and the
body pivoted from left to right and back again, grasping a
twig after each motion. Multiple bill snaps are often given
between each movement. Display Preening is carried out
often by several birds at the same time even though out of
sight of one another. This action resembles true preening
except that the bill is not closed on the wing-feathers that
appear to be treated.
Copulation occurs at the nest. The female holds her wings
open and the male hooks his feet above her shoulders. The
male clatters his bill loudly and shakes his head vigorously,
beating the female's bill back and forth with his own.
Eggs are dull white, sometimes streaked with brown; 2-5
but most commonly 3 or 4 are laid (S. Ali and S.D. Ripley
1968:94). In 90 clutches over a 6 year period at Delhi, the
clutch-size varied from 1 to 4, with an average of 2.49/clutch
(Desai et al. 1977). The eggs are laid 2-3 days, occasionally
4 days, apart; incubation starts with-the first egg and is
shared by both birds. At Delhi, for 37 eggs over a 4-year
period, the time taken to hatch was 28-32 days, with an
average of 30.2 days (Desai et al. 1977). At Bharatpur, 6
eggs in 3 different nests had a 27- to 29-day incubation
period (Breeden and Breeden 1982).
Parents regurgitate food onto the floor of the nest and,
when they are strong enough, the chicks pick it up. Small
nestings are frequently shaded from the hot sun by a parent
in the 'delta-wing' posture, with wings partially opened to
the wrist (Kahl 197Ig).
Young are seldom, if ever, left unattended until they are
about 3 weeks old (Desai et al. 1974). Predation by the ever
present house crows (Corvus splendent) and Pariah Kites

(Milvus migrant) account for heavy losses. Lowther
(1949:140) reported heavy predation on nestlings by a Pal-
las's Fishing Eagle (Haliaeetus leucoryphus) in the Mainpuri
District, India. Young birds begin to exercise their wings
after 25-30 days, but remain unable to fly until about 60-
70 days. They are usually fully independent by 85 days, but
sometimes they will return to the nest up to 115 days,
feeding largely on their own but still soliciting food from
their parents (Shah and Desai 1975b, Desai et al. 1977).
Success in rearing young is dependent on food availability,
and this in turn depends on fish stocks and the water level.
Over six seasons in Delhi, Desai et al. (1977) found that
68% of the eggs hatched and 64% of the young fledged,
giving an overall success rate of 44% or 1.1 chicks/nest.
Maturity is slow, with the plumage changing over 2 years.
By about 16 months of age, young loose the ability to vocal-
ize, and the adult plumage is attained at 3 years. Three
marked individuals first bred at 4 years old (Shah and Desai
1975b).
TAXONOMY
Formerly classified under the genus Ibis, the Painted Stork is
now combined with the other wood storks in the genus
Mycteria. The inaccurate common name of 'Wood Ibis' is,
fortunately, rarely used today.
Even though apparently not closely related, Painted
Storks have hybridized with Lesser Adjutant Storks (Leptop-
tilos javanicus) on several occasions. A male Painted Stork
and a female Lesser Adjutant Stork mated and reared young
several times in the Colombo Zoo, Sri Lanka (W.G.O. Hill
1943, A.P. Gray 1958:5), and another pair (sexes undeter-
mined) did the same in the Kuala Lumpur Zoo, Malaysia,
in 1974 (P. Olney pers. comm. 1974).
Painted Stork 57
CONSERVATION
The Painted Stork is one of the most widespread and least
threatened storks of Asia (Luthin 1987), but its future sur-
vival depends almost entirely on human behaviour and
conservation of its wetland habitat. Human behaviour, in
turn, depends very often on religious attitudes. In India and
Sri Lanka, the human population lives very much in har-
mony with wildlife, and in most areas active support is given
to birds in traditional colonies. Indeed villagers take pride
in 'their' birds. Elsewhere throughout its range, such as
Bangladesh, Burma and Thailand, this stork has suffered
not only from drainage of swamps and marshlands and the
felling of nest-trees, but also from increasingly active hunt-
ing. Young and eggs are taken from nests both for food, and
for profit. A good price is obtained by hunters, often pro-
fessionals, for young birds to be kept as pets or sent to zoos.
Increases in the human population throughout Asia, and
especially in Southeast Asia, has shown an upsurge in this
activity. Unless these and other large wetland birds are
given protection by laws that can be enforced, the Painted
Stork will become extirpated over much of its range. All
nesting colonies require protection from disturbance.
Because captive populations exist in Asian zoos (R.E.
Johnson et al. 1987a), these can be used for stocking other
zoos and for restocking wild populations, once protection is
afforded. If left undisturbed, the Painted Stork can become
fairly tame and nest in close proximity to people, even in
their villages.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 287.
 Asian
Openbill Stork
Anastomus oscitans (Boddaert)

Ardea oscitans Boddaert, Table des planches enlumineez d'histoire naturelle, 1783, p. 55: Pondicherry

Other names:

Shell-Ibis, Shell-eater, White Openbill (English); Le bee ouvert (French); Weiss-KlafTschnabel

(German); Naththai-kuththi-narai, Karu-narai (Tamil/Sri Lanka); Bellan-kokka, Beli-kava
(Sinhalese); Karu-tsoke (Burmese); Nok Pak Hang (Thai)

IDENTIFICATION
The Asian Openbill Stork is a small pale grey to whitish stork, standing about 68 cm tall, having
scapulars, flight feathers, some coverts in the wing, and tail black. Together with the African species, it
is distinguished by its peculiarly shaped bill that has a narrow gap between the two mandibles, formed
by the lower being curved downwards, then upwards, meeting the nearly straight upper mandible again
at the tip.
Adult plumage is predominantly white, with black in the wings and tail. The bill is a dull greenish-
horny colour, mottled and streaked with black and reddish; the average gap between the mandibles in
14 adult specimens was 5.87 mm (Kahl 1971c). The iris varies from dark grey to brown, and the lores
are blackish to dull blue. The legs and feet are dull flesh colour. Sexes are alike, but the male is
marginally larger and its bill is longer and heavier.
Before the onset of breeding, probably through a moult, the greyish plumage becomes immaculate
white and the black feathers take on an iridescent purplish green. The white changes back to grey soon
after the eggs are laid, in a process not well understood, presumably through feather wear and/or soiling
(Kahl 1970). [The Oriental Crested Ibis undergoes a similar colour change from white to grey, which is
apparently caused by the bird rubbing a 'black substance' from the skin of the face onto the feathers
(Uchida 1970).] There is a band of bright orange skin, about 2.0 cm wide and 15.0-17.5 cm long, on the
underside of the forearm (radius/ulna), which shows when the wing is opened (pers. obs. M.P.K.). The
bill is basically horn-coloured and blacker at the base, but becomes mixed with shiny patches of reddish
brown. The dull flesh-coloured legs and feet flush a deeper pink at the start of the breeding season.
Small chicks have sparse, pale greyish down, tinged with brown, over blackish skin. Their gular skin
is a bright pinkish purple (pers. obs. M.P.K.). Larger nestlings are a dirty grey, with black flight
feathers in the wing and tail. Immature birds are a darker grey than adults and the head and neck are
browner; they have a blackish brown mantle, wings and tail. The legs and feet are pale horn colour
(often streaked with white droppings). The short bill is dark and has little or no gap until the bird is
approximately 4-6 months old (Legge 1880:1103). In captive birds the gap continued to increase over
several years, even though the birds had no access to shelled molluscs on which to feed (Kahl 1971c,
H.G. Klos pers. comm. 1970).
During courtship displays, birds utter a series of hollow, slightly nasal honking sounds (Kahl 1972d).
This call has also been described as a 'mournful "Hoo-hoo"' (Loke in J. Huxley 1962).
60 Asian Openbill Stork
Normal flight is buoyant, with several flaps followed by a
short glide; the flapping rate is 208/min (n = 44) (Kahl
1971b). Between breeding colony and feeding grounds,
Asian Openbills soar on thermals and glide for long dis-
tances. When returning to the colony, birds often dive
almost vertically at dizzying speeds, banking, slide-slipping,
and gyrating violently (Krishnan 1960, S. Ali and S.D.
Ripley 1968:96).
The grey plumage and gaped bill are quite distinctive.
The Asian Openbill is much smaller than the Painted Stork,
with which it often nests. Its range does not overlap with the
darker African Openbill. At a distance, it might possibly be
mistaken for the larger, whiter White Stork, which occurs
within the range during migration.
DISTRIBUTION AND POPULATION
The Asian Openbill Stork historically was widespread
throughout tropical Asia, from the Indian subcontinent
(India, Sri Lanka, Pakistan, Nepalese terai) through Thai-
land to Vietnam. Populations now occur rather patchily in
suitable protected habitat (see Luthin 1987).
It no longer breeds in Pakistan and, indeed, is only a rare
visitor there (T. Roberts pers. comm.). Indian colonies are
scattered, whereas several colonies occur in protected sites
in Sri Lanka, such as in Wilpattu, Yala and Kalametiya
National Parks. The once extensive Bangladesh population
persists only along the coast in the Sundarbans and in
Ghittagong. In Thailand, one large colony and several
smaller ones are known. In fact, the largest extant colony is
at Wat Phai Lorn, near Pathum Thani, about 40 km north
of Bangkok. Other recently colonized Thai sites are at Wat
Ampuwararam, Wat Tarn En and Ban Tha Sadet (Suphan-
buri Province); the former was undoubtedly established by
a break off from the Wat Phai Lorn colony. The Asian
Openbill is now rare in Vietnam, being known from only
two sites. Information from Kampuchea is lacking.
Stragglers occur in the Malay Peninsula, south to southern
Thailand or beyond.
On the Indian subcontinent, the species moves in synch-
rony with the varying distribution of seasonal monsoons.
Henry (1971:384) reported seeing a flock of c. 100 entering
Sri Lanka, at the Jaffna Peninsula, as if arriving from India.
We have found that the Asian Openbill does not return to
its breeding sites in Bharatpur, India, when the monsoons
fail. At various lighthouses along the coasts of India, birds
dash themselves against the dome of the lantern, suggesting
that movement is sometimes at night (S. Ali and S.D. Ripley
1968:96).
It is well documented that in Thailand, the Asian Open-
bill has a well marked migration route (McClure 1974:93).
It arrives at Wat Phai Lom in October and November and
disperses in April and May after nesting, except for a few
hundred birds that remain as residents in the wet season.
Some birds travel only to northern Thailand and a few to
Kampuchea. But many migrate to the deltas of the Brahma-
putra and Ganges rivers in Bangladesh, 1500km away.
They can be seen along this route in large numbers; and,
outside the regular periods of migration, the species occurs
in small numbers at almost any place along the well defined
route. This amazing migratory pattern was discovered dur-
ing the extensive ringing programme in 1968 to 1970, under
the direction of Dr H. Elliott McClure.
Although reduced in geographic extent and overall num-
bers, the Asian Openbill remains the most abundant Asian
stork (Luthin 1987). Its largest colony at the Wat Phai Lom
Buddhist temple in Thailand numbers between 8000 and
14000 nests in a relatively small area (4.7 ha). In 1980,
about 30 000 birds were counted including young, and the
other Thai populations totaled another 200-1000 indi-
viduals. During the period of December 1988 to February
1989, a total of 20 358 Openbills were counted in India and
an additional 4172 in Sri Lanka (D.A. Scott and P.M. Rose
1989:56). In the 1950s, the species was common throughout
Bangladesh, but is now reduced to about 30 in the Ghitta-
gong Hills and 50-70 in the Sundarbans (M.A.R. Khan
1984, Luthin 1984a). These areas continue to serve as the
off-season home for many of the Wat Phai Lom birds from
Thailand.
ECOLOGY
The Asian Openbill Stork is found in marshes, jheels,
flooded rice-paddies, mangrove swamps, and less commonly
along river banks or on tidal mudflats. Loss of natural
habitat in Thailand has meant that this species has shifted
to feeding in rice fields and the edges of canals and rivers,
where apple-snails (Pila) are still plentiful.
Commonly foraging is in small to medium-sized flocks.
Feeding is by tactile or visual methods (Kahl 1971c). Snails
are secured by the forcep-like bill, while the bird Walks
Slowly or Stands in shallow water. The Probing bill is
jabbed downward, and is quickly closed on any prey. The
snail is extracted from its shell in exactly the same way as by
the African Openbill, using the tip of the thin lower man-
dible to extract the snail body, leaving the relatively intact
shell behind (Kahl 1971c). The common supposition (J.S.
Huxley 1960) that the unique bill is regularly employed in
the manner of a 'nut-cracker' does not seem to apply,

especially in the case of the favourite food, large Pila snails.
However, small snails are sometimes crushed between the
mandibles (Henry 1955, Mukherjee 1974).
Jerdon (1864:766) reported that blinded captives
removed a snail from its shell without crushing it, and he
gave a graphic description of the extraction process:
The bird secured a shell by its feet, and after sundry
alterations of its position, succeeded in cutting off the
operculum as cleanly as if it had been done by a razor,
but so rapidly that I was unable to see the exact way in
which it was accomplished. It then inserted the tip of
its clumsy beak into the open mouth of the shell, and
after working it about for a short time, pulled out the
entire shell-fish almost to its utmost tip.
Many individuals have the extreme tip of their lower
mandible curved slightly to their right (N.F.R. Snyder and
H.A. Snyder 1969). This asymmetry may help the bird
sever the columellar muscle of the apple-snail. Such asym-
metry is not usually found in the African Openbill (Kahl
1971c), and it would be interesting to learn if this is because
the Asian birds feed more regularly on snails that coil in a
given direction, as suggested by N.F.R. Snyder and H.A.
Snyder (1969).
Although foraging is usually done in water, in Thailand
Asian Openbills feeding nestlings during the peak of the
dry-season sometimes follow tractors in dry fields and ob-
tain aestivating snails turned up by the blade (P. Round et
al. unpub. report).
E.C.S. Baker (1929:334) describes the main diet as being
molluscs, chiefly those of the large Pila snails, but the Asian
Openbill occasionally eats bivalve molluscs, crabs, frogs,
insects and other small animals found in its accustomed
feeding marshes (S. Ali and S.D. Ripley 1968, McClure and
Kwanyuen 1973, Breeden and Breeden 1982). In the Sun-
darban area of northeastern India, Mukherjee (1974) col-
lected 72 adult Openbills for stomach analyses. He found
that 85% of the wet-season diet consisted of molluscs
(mainly Pila snails), but that in the dry season more crus-
taceans, fish, reptiles, insects, etc., were taken. An adult
Openbill, collected at Bharatpur, India, in September 1967,
contained over 150 small snail bodies (Kahl 1971c).
In the Indian region, during normal rainfall seasons,
Asian Openbills arrive in an area at the commencement of
the rains, coinciding with the snails' re-emergence from dry-
season torpor. In Thailand, birds arrive and breeding starts
at the beginning of the dry season (P. Round pers. comm.
1990).
At Bharapur, India, we found that in 1987, when no rain
fell, the Asian Openbills did not appear at all, whereas in
1988, exceptionally heavy rains brought the storks back and
they commenced nesting at their traditional sites. However,
this nesting failed because of excessively high water levels,
owing to supplementary water pumped from a nearby canal
and river system. This additional water raised the levels,
apparently too high for the birds to forage, leading to aban-
donment of the traditional site and movement to a new site.
The disruption of traditional feeding areas caused the storks
to leave their nests unattended, to feed. Their eggs were
Asian Openbill Stork 61
then robbed by House and Jungle Crows (Corvus splendens
and macrorhynchos) as well as by Pariah Kites (Milvus
migrans), which had prospered in the drought year and
whose numbers were correspondingly high. Additionally
these delays and disruptions to the normal breeding pattern
meant that Painted Storks had started to nest before the
Asian Openbills were fully established, leading to heavy
fighting between the species, and the resultant desertion by
many of the much smaller Asian Openbills. We saw large
parties of these birds feeding in small ponds and flooded
fields well away from the colony, many still retaining bright
courtship colours, including red legs. Thus nesting failed in
two successive years, one because of drought and one be-
cause of excess water, exacerbated by artificial flooding.
Thus it seems that many of the ecological requirements of
the Asian Openbill Stork revolve around having suitable
water supplies on its feeding areas.
BREEDING
The species is highly social, its colonies being exceptionally
dense, with as many as 62 nests in a single tree in India
(Hume and Oates 1890:225) and 156 in a single Sterculia
foetida in Thailand (McClure and Kwanyuen 1973). In
India, this stork often breeds in large mixed colonies with
other storks, ibises, spoonbills, herons and cormorants.
However, the largest colony, in Thailand, contains only
Asian Openbills. In India and Sri Lanka, they are usually
the first wading birds to begin nesting at the beginning of
the monsoon. In Thailand, however, nesting begins at the
end of the rainy season and progresses throughout the driest
time of year.
Nest-platforms are placed in the tops of trees, often over
water. In India, they sometimes use large trees, such as
peepul (Ficus religiosa) or neem (Melia azadirachta) (Hume
and Oates 1890:224), or smaller trees, such as babul (Acacia
arabica] (S. Ali 1953:533). In Thailand, we have seen up to
10 nests on a single bamboo shoot. They were precariously
sited and the close proximity of nests can lead to inter-
ference from neighbours. This over-use of all available sites
is due to the need for a safe nesting area, provided only on
the temple grounds where the birds are safe from human
persecution (see Conservation). Bamboo nesting sites are
subject to wind and storm loss and, as a result, productivity
is low.
Courtship behaviour is described in detail in Kahl
(1972d). Unpaired males choose display sites, while females
move around observing their displays. In defence of its site,
a bird will use Forward Clattering, very subtle Bill Snaps,
and Arching Displays. Sexual displays include Swaying
Twig-Grasping, Advertising Sway and the Up-Down. The
latter greeting display is often given by both members of the
pair, especially after the return of one to the nest. In this
display, the bird stands erect, arching its neck forward and
pointing its bill horizontally or slightly upward. It then
moves its head slowly downward and utters a series of nasal
honking sounds. The Up-Down often grades into Swaying
Twig-Grasping, and is preceded and followed by Bill Snaps.
During Copulation Clattering, the clattering sound comes
62 Asian Openbill Stork
from the male vigorously slapping his bill against that of the
female. Pair-formation and behaviour during breeding are
generally similar in the Asian and the African species.
However, in hostile situations, the African birds erect the
feathers of the crown and occiput, whereas the Asian birds
are less prone to do so. Head shaking and quick rubs of the
crown on the shoulders and upper back are also less com-
mon in the Asian birds. The Advertising Sway has not yet
been reported for the African Openbill. The displays of both
species are somewhat less ritualized than in some other
storks (Kahl 197If), though the Up-Down greeting displays
of the openbills are quite similar to those of the wood storks.
Both sexes cooperate in the building of the nest, with the
male collecting most of the sticks. Birds make up to 30 trips
per day with material, and nests may take as long as 11 days
to build (Mukhopadhyay 1980). Five nests measured 100-
125 cm in diameter (Mukhopadhyay 1980) and were built of
small sticks and lined with leaves and other vegetation.
Fresh foliage is brought to add to the nest even during
incubation, possibly to provide moisture.
Between 2 and 4 eggs are laid, rarely 5. They are dull
white and are broad oval in shape, slightly compressed at
the smaller end (E.C.S. Baker 1929:334). Both birds share
incubation. Estimates of the incubation period vary from
24-25 days (S..A11 and S.D. Ripley 1968:97), 27-29 days
(McGlure and Kwanyuen 1973), to 28-30 days (Mukho-
padhyay 1980); we believe the longer periods (27-30 days)
to be more accurate. Regurgitation of water over eggs was
noted in India (Kahl 1970, Mukhopadhyay 1980) but it has
not been reported in Thailand. On rare occasions, 3 adults
have been observed tending a nest (B. Amget pers. comm.,
pers. obs. M.P.K.).
Small chicks are shielded from the sun by the parents
standing on the edge of the nest covering them with open
and drooping wings (often not as erect as Painted Storks).
Nestlings are showered with water by their parents during
the hot part of the day (Breeden and Breeden 1982, B.
Amget pers. comm.). Both parents feed the young by regur-
gitation into the nest, but some chicks will grab at the
adult's bill and catch the food before it reaches the floor of
the nest. Mukhopadhyay (1980) reported that chicks were
fed 5-6 times per day.
Predation on eggs and young in India is by monitor
lizards (Varanus sp.). Jungle Crows (Corvus macrorhynchos]
and several species of birds of prey. Breeding success at a
colony in northeastern India was about 50% (Mukhopad-
hyay 1980). At the large Wat Phai Lorn colony, in Thailand,
usually only 2 young survive per nest; in years without wind
storms, the survival is higher. Predation is neglible there,
but the nestlings are victims of parasitism by various ticks,
flies and mites (McClure and Kwanyuen 1973).
First flights are at about 35-40 days, and the young
finally leave the nest at about 6-8 weeks (McGlure and
Kwanyuen 1973, Mukhopadhyay 1980). At Bharatpur,
India, fledged young are active in the nesting areas often
before other species, including Painted Storks, have finished
incubation. When really ready to leave the nest, the young
flap their wings frequently, and when fully fledged, they
gather in groups away from the nest.
TAXONOMY
The Asian Openbill is very closely related to the African
Openbill Stork, from which it mainly differs in colour. The
two openbills form a natural group, here designated as a
tribe, with the four wood storks of the genus Mycteria. Simi-
larities in many of their courtship displays indicate this close
relationship (Kahl 197If, 1972b, 1972d, 1972e).
CONSERVATION
Toleration by humans of nesting waterbirds in India and
protection of temple sites in Thailand have served the
species well. However this tolerance is not matched in
Pakistan, where it now no longer breeds because of con-
tinual persecution, or in Bangladesh, where a tradition of
hunting goes back to the British. Trapping remains a prob-
lem there. In Thailand, protection is given by Buddhist
monks in their temple grounds at Wat Phai Lorn, but the
large population of birds and the accumulated excreta have
killed a number of the trees and are threatening the others.
Inkapatanakul (1986) found that the heavy use of bamboo
was due to the loss of other trees. It has even been suggested
(Sitwell 1984) that the species of bamboo used by Asian
Openbills at Wat Phai Lorn (Bambusa arundinaced) is due for
a major die-off about 1993, which could be disastrous for the
storks nesting there.
Drainage of natural marshes in Thailand has led to the
birds seeking food in cultivated fields, and the danger from
pesticides has increased. The long flight path on migration,
over what has become hostile territory, results in heavy
losses particularly of first-year birds. Whilst Indian and Sri
Lankan birds are afforded unmolested breeding areas, they
are subject to drought in some years, artificial flooding in
others, and a lack of alternative colony sites. Other breeding
populations in Southeast Asia are poorly documented, but
are certainly in trouble. Various evidence from the biology
of the species demonstrates its dependence on particular
hydrological conditions. With both drought and extreme
artificial flooding causing nesting failure, water manage-
ment in feeding sites becomes critical.
The Asian Openbill Stork is in fact a species that could
benefit from active management throughout its range. Tem-
ple sites are particularly suitable for active management.
Remnant colonies could provide stock for repopulation, if
productivity were increased and additional nesting and
feeding sites provided. All colony sites need to be provided
complete protection from both poaching and disturbance
(Luthin 1987). Nesting Asian Openbills are in competition
for trees with wood cutters; additional sites can be provided
by protecting tree stands, and present sites can be enhanced
through tree planting and construction of artificial nest sites
(Inkapatanakul 1986). Formerly, river flooding washed
excrement away annually; now flood control has prevented
this and the birds' excrement accumulates. In restricted
areas where nutrient accumulation is killing trees, some
dilution of nutrients may be feasible.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 288.
 African
Openbill Stork
Anastomus lamelligerus Temminck

Subspecies:
Anastomus lamelligerus lamelligerus Temminck
Anastomus lamelligerus Temminck, 1823, Planches Coloriees, livr. 40, pi. 236: Senegal
Anastomus lamelligerus madagascariensis Milne-Edwards
Anastomus madagascariensis Milne-Edwards, 1880, Compt. Rend. Acad. Sci., Paris, 91, p. 1037:
Madagascar

Other names:
Claw-billed Stork (English); Bec-ouvert africain (French); Schwarz-Klaffschnabel (German);
Masoklo (Luo); Etongorokofu (Kwangali); Oopbek-ooievaar (Afrikaans); Famakacora (Malagasy)

IDENTIFICATION
The African Openbill Stork, a medium-sized stork, standing 55-60 cm, is an all black bird with a stout,
distinctively shaped bill, which is open for part of its length. The peculiarly shaped bill, from which it
gets its generic and common names, is a brownish horn colour but paler, often a dull yellowish tan,
basally. The upper mandible is essentially straight, but the lower is curved upward, leaving a gap
between the two, which meet again at the tip; the average gap in 16 adult specimens was 5.9 mm (Kahl
1971c). The lores and the skin around the eye are dull-bluish to blackish. The legs and feet are black
and the eye is brown. The iris is dusky brown. The sexes are alike, but the male is on average larger,
1250 g compared with 1000 g for the female, and the male's bill is larger.
The dark plumage is enlivened during the breeding season with shiny, sequin-like feathers of
iridescent metallic brown, black, green and purple, most pronounced on the abdomen, owing to highly
modified feather morphology (Stettenheim 1976). Those on the breast and abdomen are curly and jet-
black, whereas those on the back and wing-coverts are flattened and green, brown or bronze
(Mackworth-Praed and Grant 1973:67, pers. obs. M.P.K.).
Newly hatched chicks weigh 35-40 g (n = 6) and are covered with a sparse, wispy grey down on the
back, wings, and head. The skin of the head and neck are pink and that of the body 'almost purple'
(Anthony and Sherry 1980). The bill is black, with the distal third cream-white, and becomes mostly
black by about 25 days. The legs and feet of small nestlings are pinkish-orange, later fading to fleshy-
pink and then dingy grey. The iris is dusky brown. At about 10 days a dense, sooty black down emerges;
at this stage the naked gular pouch is a bright pink (pers. obs. M.P.K.).
Immature birds are duller with brown feather-edgings below and on the back and upperwing-coverts,
and the back of the neck speckled white. The bill is blacker, lacks the paler base and initially is shorter
and lacks the gap of the adult (Anthony and Sherry 1980). Typical bill configuration develops slowly,
possibly taking several months.
The African Openbill is usually silent, except at the breeding colony. Courting adults utter loud,
sonorous, rather raucous croaks or honks, 'Horrrh-horrrh', audible at up to 100 m. During copulation, a

clattering sound is produced by the male beating his bill
against that of the female. The Begging call of nestlings is
similar to the Yellowbilled Stork, but higher-pitched and
more nasal (Brown et al. 1982:176, pers. obs. M.P.K.).
In flight the tail appears broad and rounded and the feet
extend beyond it (Bannerman 1930:103). Often birds soar
on thermals, especially during long-distance movements
(Ruwet 1963). In flapping flight, a series of flaps alternates
with a short glide; the flapping rate averages 201/min
( » = 18) (KahI1971b).
Its gaped bill makes the African Openbill unique and
easily distinguishable from other species. At distances too
great to discern the bill, it could be mistaken for other dark
waterbirds, such as the Glossy Ibis or Black Heron (Egretta
ardesiaca). It is geographically isolated from the lighter-
plumaged Asian Openbill Stork.
DISTRIBUTION AND POPULATION
This wide-ranging stork is found in suitable habitat
throughout the African tropics south of the Sahara. Most of
the breeding seems to take place in the Southern Hemi-
sphere. It mainly occurs below 1500 m elevation.
In West Africa, there is a report of breeding only from
Sierra Leone (Bannerman 1953:175). It has nested on occa-
sions and in small numbers in Chad and Ethiopia (Salvan
1967, SchaufFele and Schiitz [sic] 1968). In East Africa, it is
widespread through Kenya and Uganda, breeding errati-
cally at Garsen, Kisumu, Rwenzori National Park, Busoga
and Jinja. In Tanzania it breeds in large numbers at Utenge
and Wembere and in smaller numbers elsewhere (Britton
1980:20). Breeding colonies have been found throughout
Zambia in all months of the year. In southern Africa, this
species is uncommon south of the Zambezi (Clancey 1964a)
and breeds in Zimbabwe, northeastern Botswana, northern
Namibia, and northern South Africa (Dutton 1972, Braine
African Openbill Stork 65
1974, Newman and English 1975, Tree 1978, Anthony and
Sherry 1980),
In Madagascar there is a separate race, A.I. madagascarien-
sis, distributed in the western parts of the island, including
the highlands (Bonvallot and Randrianasolo 1975, Lan-
grand 1990).
The African Openbill wanders considerably within the
wetter parts of the African tropics, probably moving back
and forth across the equator. Its movements are not well
known, but it becomes locally abundant when conditions
are right. It is primarily a non-breeding visitor north of the
equator and is more common in the east than in the west. It
arrives in the dry season, presumably from breeding areas in
the south. Large flocks gather at Lake Turkana, Kenya,
each November-December, suggesting well-established
movements from breeding sites in the Southern Hemisphere
(Britton 1980:20). We have seen it in a mixed colony on the
Tana river in January 1983, when it took advantage of
exceptionally heavy short rains to breed, but in other years
it was absent in the same area, when the short rains failed
(Hancock 1984:41).
The population is probably stable (Brown et al.
1982:176). The African Openbill is locally numerous year-
round in East Africa, especially near breeding and mi-
gratory sites, and may be extending its range there; flocks
up to 7000 have been reported (Kasoma and Pomeroy
1987). It is the most numerous stork for part of the year
(Dec-May) in Sudan but is uncommon in the highlands of
Kenya. In Tanzania, breeding colonies can number in the
thousands. The largest reported colony there was of 5000 or
more nests (Kahl 1968), but this may have been too high an
estimate (Stronach 1968). O. Langrand (pers. comm. 1989)
reports it to be common in Madagascar, between Maha-
janga and Morombe, particularly near Bemamba Lake and
Morondava where groups of 15-20 birds are commonly
seen. Conversely it no longer is seen at Alaotra Lake.
ECOLOGY
Characteristically a floodplain species, the African Openbill
Stork is found mostly in freshwater wetlands, and not
usually in heavily forested areas. It requires shallow water,
such as ponds, streams, rivers and temporarily flooded
marshes; it will sometimes also forage on drier ground if
appropriate food is present. In West Africa, it is occasion-
ally seen on coastal mudflats (Bannerman 1953:175). Its
main habitats in Madagascar are shallow lakes, rice pad-
dies, and occasionally marine coasts and mangrove swamps
(Langrand 1990). At other times it travels in large flocks
and can be found in any habitat, including ploughed fields
(Britton 1980:20).
This species feeds either alone or in groups, or with
Sacred and Hadada Ibises (FJ. Jackson 1938:74). The
foraging strategy involves both tactile and visual feeding.
Both day and night feeding have been reported (Clancey
1964a:46). Birds will probe deeply with bill open, often with
head submerged, to capture snails in ponds and shallow
lakes, which are often covered by floating vegetation. They
may Stand in one place repeatedly Probing before moving to
66 African Openbill Stork
a new spot. We have seen up to 50 birds feeding close
together in one such lily pond in East Africa (Hancock
1984) and riding on the backs of hippopotamuses to catch
prey disturbed by these animals (Kahl 1971c). When feed-
ing in groups, mutual harassment and attempted prey-
robbing have been reported (Niven and Niven 1966a).
Early authors suggested that the unique gap in the bill of
this species was used to carry and then crush the shell of the
snail (Chapin 1932a:472, F.J.Jackson 1938:74, J.S. Huxley
1960). We have found, however, that the snail is most often
held in the tip of the bill, and the extraction accomplished
by holding the shell against the substrate with the distal end
of the upper mandible. The tip of the knife-thin lower
mandible is then forced under the operculum, severing it
from the body. The stork then shakes its head to release the
body of the snail from the shell, which remains essentially
intact. This process can take as little as 15 s (Kahl 1971c).
Mussels are opened in a similar manner or by holding and
squeezing until the animal opens its shell (Root 1963). In
the latter case, when a mussel is caught, it is carried to the
shore where piles of discarded shells can accumulate as the
same site is used over and over again (Kasoma and
Pomeroy 1987). Such specialized actions are aided by
unique quadrate bones in the skull which have one narrow
condyle and assist this form of feeding. Comparison between
the bill shapes of openbills and other snail-eating species is
discussed by J.S. Huxley (1960). Because openbills must
spend some time manipulating their prey before swallowing
it, other species, such as gulls, may sometimes attempt to
steal it (Steyn 1988).
The African Openbill Stork is a highly specialized species
that depends almost entirely on molluscs. Heuglin (1869-
73:1121) considered the overall diet to be broader, including
fish, frogs, crabs, locusts, beetles, worms, mussels and
snails. Since then, most other observers have mentioned
only snails and freshwater mussels, including the genera
Lanistes, Limicolaria, Achatina, Pila and Mutela (references in
Kahl 1971c, Anthony and Sherry 1980). While some of the
items of food described by Heuglin may be occasionally
taken, it is clear that this stork feeds almost exclusively on
mussels and snails, both from fresh water and dry land, with
snails of the genus Pila predominating. It is the emergence
of these snails, brought about by the onset of the rains, that
triggers the arrival of these birds in their breeding area.
Apparently the African species eats more mussels than does
the Asian species (J.S. Huxley 1960, 1962), but they are not
always successful in opening all the mussels that they catch
(Root 1963). Many hours of observation have been spent
without seeing these birds catch anything except molluscs
(Bell-Gross 1974, pers. obs. M.P.K.).
Captive-reared nestlings strongly favoured a diet of mol-
lusc bodies, and refused other food (e.g. small fish) even
when ravenously hungry (Kahl 1972f). A captive 15-day-
old nestling weighing 148g consumed 143g of snail bodies
(nearly 97% of body weight) in one 12 h period. At 37 days
of age, a nestling weighing 720 g ate 243 g of snail bodies in
15-20 s and still begged for more (Kahl 1971c).
The African Openbill is a very gregarious stork, in its
breeding, foraging, flying and roosting. Often they arrive in
a large flock and then disperse to feed individually. Very
heavy rains, with the resultant flooding and abundant
supply of snails, often result in African Openbills suddenly
appearing in an area and starting to breed. In years of
exceptional flooding, breeding may span long periods
(Anthony and Sherry 1980).
It has been reported that this species is sometimes used as
an animated feeding-perch by Carmine B.ee-eaters (Merops
nubicus) (Cunningham-Van Someren 1970).
BREEDING
African Openbill Storks breed colonially, often alongside
ibises, spoonbills, herons, darters, cormorants and other
storks. They are opportunistic breeders, and nesting occurs
only in years when local food supplies are plentiful; it does
not occur regularly in the same place. Nesting is in trees or
bushes, often over water, and sometimes as many as two
dozen or more nests of this species are found in a single tree.
Most populations of this species nest in the rains, at the
height of flood-plains inundation. Commencement of breed-
ing is probably related to the emergence from aestivation of
snails (Pila).
Males stand at the nest site and are approached by
unmated females. Courtship and pair-formation are basi-
cally similar to the processes in other storks (Kahl 1972d).
Males secure nest sites and perform a variety of displays.
One prominent display in the courtship of the Asian Open-
bill is the Advertising Sway; this display has not been con-
firmed in the African species —contrary to the statement in
Brown et al. (1982:177)—but probably does occur. Com-
monly given is the Up-Down display, which involves an
arching of the neck and gaping of the bill; raucous calls are
uttered but no bill snaps are given. The Swaying Twig-
Grasping display is often performed. The Bill Snap is fre-
quently given early in pair-formation but less after pairing is
completed. Forward Clattering is given by males defending
sites from other males, and by females threatening other
females. Disturbance near the nest by animals or humans
elicits the Arching display, and this serves to warn other
birds of danger. Copulation Clattering is accompanied by
the male shaking his head vigorously and striking his bill
against the bill of the female; he does not clatter his own bill,
as most other species of storks do.
Nest platforms are made of twigs and lined with grass,
leaves, sedge, various water plants; the inner lining is often
damp. Nests measure 45-75 cm in outside diameter and
about 20cm across the inner cup (Rand 1936:341, Heiden
1973, Anthony and Sherry 1980). They are built either by
repairing an old nest or building afresh with sticks and
reeds, and lining with weed collected from the bottom of a
pond. We have seen this carried out by birds bringing quite
large dripping piles of weed, even after incubation has
started, and the nest is kept wet throughout, sometimes by
regurgitating water. Loud calls are given by both birds at
the nest during these actions. Initial nest building or repair
spans about a week before egg-laying (Anthony and Sherry
1980).
Eggs are laid at 2-day intervals; 28 clutches varied from

3-5 eggs, averaging 3.9 (Maclean 1985:68). They are dull
white and become heavily stained.
Both birds incubate, often with the partner standing
nearby. Incubation has been estimated as 25-30 days
(Brown et al. 1982:177). Egg predation has been reported
by a Gymnogene or African Harrier Hawk (Polyboroides
typus) from Zimbabwe (Anthony and Sherry 1980).
Nestlings have shorter, straight bills, but possess a wide
buccal cavity, which enables a nestling weighing 45 g to
swallow large snails up to 5 g. Begging is done with open
wings flapping and slightly nasal cries. Often the bill is
pointed upwards exposing the pink gular skin of the chick.
We found that captive nestlings usually refused food other
than mollusc bodies even when ravenously hungry (Kahl
1971c). Parental feeding of young is accomplished by regur-
gitating food on to the floor of the nest. Parents often
regurgitate water over the nestlings.
Flight feathers begin to appear at about 25 days, and the
body is fully covered with contour feathers by 40 days. The
fledgling period is estimated to be 50-55 days (Brown et al.
1982:177, pers. obs. M.P.K.); that of 80 days, given by
Anthony and Sherry (1980) and Maclean (1985:68), is
almost certainly too long. Nesting success varies greatly
with feeding conditions, from complete failure to 2-A young/
nest (Brown et al. 1982:177).
TAXONOMY
The two openbill species are closely related to each other,
morphologically and behaviourally. The Up-Down and
Copulation Clattering displays show close resemblance to
homologous displays seen in the several species of wood
storks (Mycteria). The most significant morphological differ-
ences between the openbills and wood storks involve the
cranium and bill (Beddard 1901, Verheyen 1959, Kahl
African Openbill Stork 67
1971c). Because of their similarities to each other, and
differences from the other storks, we have combined the
wood storks and openbills in the tribe Mycteriini (Kahl
1972e).
The separation between the two subspecies is based on
the Madagascar bird having more pronounced longitudinal
ridges on the bills of adults and more white spots on the
necks of juveniles (Bangs 1918, Chapin 1932a:470, Banner-
man 1930:103); some authors report the Madagascar bird
as larger, but the two specimens we measured in the Leiden
Museum were smaller (see Appendix). Sharpe (1898:309)
saw no reason to separate the Madagascar population. We
tend to agree, but, pending further study, have retained it
tentatively as a separate subspecies.
CONSERVATION
The African Openbill Stork is locally common and can form
flocks numbering over 7000 in some seasons. In Africa, its
population is considered to be stable, in part because its
colonies are smaller and more widely spread than are the
breeding sites of the Asian Openbill (R.E. Johnson et al.
1987a, Luthin 1987). The species may actually be expand-
ing its range in eastern Africa. In Madagascar, it was
common in the lowlands (Rand 1936), but in recent years
the Madagascar population has been reduced. Its elimin-
ation from such places as Alaotra Lake reflects the increase
in human population there, where hunting and poaching
continue to take place.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 289.
 Black Stork
Ciconia nigra (Linnaeus)

Ardea nigra (Linnaeus, 1758, Syst. Nat., ed. 10, p. 142: northern Europe ( = Sweden)

Other names:
Schwarzstorch (German); Cigogne noire (French); Zwarte ooievaar (Dutch); Svart stork (Swedish);
Sorte Stork (Danish); Ciguena negra (Spanish); Bocian czarny (Polish); Ghyorny aist (Russian);
Grootswartooievaar (Afrikaans); Endongondongo (Kwangali); Mokoroane (South Sotho); Unocofu
(Xhosa); Gandahari, Gamdayaki, Gamyaki (Hausa); Surmal (Hindi); Hei-kuan, Wu-kuan, Hei-chu-
chi (Chinese); Hyun hak (Korean); Nabe-koh, Kuro-koh (Japanese)

IDENTIFICATION
This medium-sized stork, standing 90-100 cm tall, is entirely glossy black save for the white of its lower
breast and abdomen, which extends to the long undertail-coverts that form a frilly fringe to the black
tail.
The adult's head, neck and upper wings are glossed with metallic purple and bronze, and its back
with an iridescent green. The tail-feathers are slightly duller and browner. Feathers of lower foreneck
are elongated, forming a rather shaggy neck-ruff that can be erected in displays or when the ambient
temperature is low. The bare skin around the eye, the bill and the legs are red; the iris is brown. The bill
and legs are red. The sexes are similar in plumage, although the male is larger and has a longer, heavier,
and sometimes more recurved bill (see photo in Siewert 1932b:33).
Birds moult into fresh, bright plumage in the spring (Dementiev and Gladkov 1951:444), and their
iridescence is stronger, especially on the head, neck and wing-coverts of the male (Siewert 1955:67, pers.
obs. M.P.K.). During courtship, the red skin of the face, bill and legs deepens in colour to a rich scarlet.
The area of red loral skin expands during displays and contracts when the bird is relaxed, so that the
bill and orbital-skin are separated by black feathers in non-displaying birds (pers. obs. M.P.K.). After
the breeding season, these soft parts take on a darker, less brilliant, or more brownish tint (Dementiev
and Gladkov 1951:445, Cramp 1977:328). Some sources state that a complete annual moult lasts from
May-June through winter (Naumann 1838:283, Cramp 1977:328).
Nestlings are covered with a sparse down at first, which is superseded by a thicker down later; it is
white with a greyish hue and quickly becomes soiled. Their bill is yellow, and the legs and feet are pink
or flesh-coloured; later the bill is orange-yellow at the base, grading to pale olive distally, and the legs
are pale olive. They develop full contour-feathering at about 2 months. The progression of the nestling's
development is illustrated by photographs in O. Heinroth and M. Heinroth (1924-3 l:fig. 170) and
described in more detail in Schroder and Burmeister (1974:6-7). After fledging the immature bird is
browner, without the iridescence of the adult. The neck and wings are duller, flecked, especially on the
lower neck, with lighter brown feather-tips, and some back feathers show a slight greenish tinge. The
bill and legs are pale grey-green, as is the facial skin. The bill and legs become red in the spring of their
second year, i.e. when about 10 months old (Cramp 1977:328).
The adults have more developed vocal calls than White Storks —usually a whistling or prolonged
piping or hissing—but their bill-clattering is much less developed. This could be because in the Black
Storks' more secluded nesting sites, short-range communication between mates is more important than
long-range threats against intruders. The bisyllabic calls have the sound of a sibilant inhale/exhale and
are given at a rate of about 2 couplets of whistles per second (pers. obs. M.P.K.). Their vocalizations
have been rendered as 'Cha-lie, cha lie, cha lie' (Siewert 1932a, 1955:75), or 'Huji-ji', 'Chi-chu',
70 Black Stork
Tleeeee, he fleece, hefleeeeehe'or 'Hhiio . . . hhiio' (Cramp
1977:327). A melodius flight call, *Fuo' is rarely heard
(Bauer and Glutz von Blotzheim 1966:416), but birds mi-
grating at night are heard calling (Collman and Croxall
1967). Various begging calls of the nestlings are described in
Cramp (1977:327; further details are in Schroder and Bur-
meister 1974:26-27).
Black Storks are agile fliers and able to manoeuvre nim-
bly between the branches in dense forest. The Black Stork is
said to fly with faster wing-beats than the White Stork
(Bannerman 1957:22). However, upon measurement this is
found to be not the case; in normal, level flight, the average
flapping rate of the Black Stork is 159/min (n = 22) versus
170/min for the White Stork (Kahl 1971b). Although it does
soar and glide on long-distance migration, this species does
not so readily soar on thermals as does the White Stork; it is
said that this is because the Black Stork has narrower wings
(Cramp 1977:324). On migration it usually moves in small
groups and does not often travel with flocks of White Storks;
it does often migrate with birds of prey (Porter and Willis
1968). Some migratory movement takes place at night
(Dementiev and Gladkov 1951:440, Collman and Croxall
1967). Especially near the nest, aerial acrobatics have been
reported and mild aerial hostile encounters with whistling
vocalizations occur (Fincham 1971).
The Black Stork can be mistaken at first sight for Abdim's
Stork in Africa, but the latter species is much smaller, with
white on the lower back and rump, and has greenish legs
and bill. The Black Stork can be clearly distinguished from
the Woollynecked Stork, with which it is frequently seen
during migration in northern India, by its black neck, red
bill and legs, and larger size.
DISTRIBUTION AND POPULATION
The Black Stork has the most extensive breeding range of
any stork. It does or has occurred throughout much of the
Palaearctic. It ranges from Europe—discontinuously from
Scandinavia, Germany, and Portugal—eastward through
the former USSR and Mongolia to the Maritime Territory,
and northeastern China. It also breeds in southern Africa,
from Zambia and Malawi to Cape Province.
Within this overall range, it is thinly spread and widely
dispersed and in many areas has been declining. It has been

extirpated from its former breeding grounds in southern
Sweden, Denmark, Finland, the Netherlands, and, with the
exception of a few odd pairs, from southern and western
Germany, France, Greece, and Korea (Loiseau 1977,
Luthin 1984a, 1987, Sonobe and Izawa 1987:43). Signifi-
cant nesting still occurs in Portugal, Spain, eastern Ger-
many, Hungary, Poland, Czechoslovakia, USSR, and poss-
ibly several other countries of eastern Europe.
In the USSR, the species is found as far north as along the
Baltic coast near Leningrad, Vologda and Kirov; in Siberia
on the eastern slopes of the Urals it extends north to 61°N. It
occurs patchily across much of the eastern USSR south of
60-63°N—in the mountainous and forested regions, but not
in the steppe areas—to the Pacific Ocean (further details in
Dementiev and Gladkov 1951:437, Vaurie 1965:85). It nests
in the northern portion Mongolia, down to the Gobian
Altai; and across much of northern China, westward to
Sinkiang (Etchecopar and Hue 1978:74; Meyer de Schauen-
see 1984:138). In northeastern China, it may breed (for-
merly at least) south to about the vicinity of Beijing (T.H.
Shaw 1936:124). It used to nest in southeastern Korea, near
Andong, Kyongsang-Pukto (Austin 1948:46), but has not
been reported to do so since 1966 (P. Won 1973:18); during
the mid-1980s Black Storks were observed throughout the
breeding season in northeastern Korea (Hamgyong-Pukto)
and may be nesting there (Sonobe and Izawa 1987:44).
Several Black Storks were reported in a colony of Black
Kites (Milvus migrant) and Grey Herons (Ardea cinerea) near
the Caspian coast in northern Iran during April and July,
but it is not clear whether the storks were breeding (Pass-
burg 1959). A nesting site is known from central Iran, in a
wetland near Isfahan (Luthin 1984a). Tom Roberts (pers.
comm.) considers it possible that one or two pairs might
nest in the Tobakhar mountains of Baluchistan, north-
western Pakistan, and J.A.W. Anderson (pers. comm.)
claims to have had 2 young newly fledged birds brought to
him at Quetta, Pakistan, in 1969 by hunters who found a
nest in that area. However, it is unlikely that they now
remain, as hunting pressures on breeding and migratory
birds of all species has intensified in recent years in Pakis-
tan. There are several summer sightings from central Afgha-
nistan but no definite evidence of breeding there (Paludan
1959:66).
Migration
Most Black Storks are migratory, but in a few cases popu-
lations are dispersive or sedentary. During its extensive
migrations, the species occurs throughout much of the Afri-
can and Asian tropics. The majority of eastern European
birds migrate to Africa, whilst those from western Asia
winter primarily in northern India, and those from further
east apparently winter mainly in southern China. Some of,
the Iberian population, as well as those in southern Africa,
are essentially non-migratory, although they may wander
widely in the non-breeding season.
Although mainly migrating along defined routes, the
Black Stork is apparently somewhat less dependent on soar-
ing than the White Stork. Some passage birds have been
reported from Italy, the Aegean Sea and various Mediterra-
Black Stork 71
nean islands, and they will cross wider stretches of water
than White Storks normally do (Bauer and Glutz von Blotz-
heim 1966:421, Cramp 1977:324).
Birds leave Europe, except from Iberia where they are
mainly resident, from early August to October, with the
greatest exodus in September. Nearly all European birds
travel to Africa. Relatively few travel via Gibraltar and visit
West Africa, because so few still breed in western Europe.
Moreau (1967) reported a few birds in Nigeria and Chad,
and some were seen in the Senegal Delta by Roux (1972).
There are scattered reports from several other West African
countries (Bouet 1955a). There is a small but regular spring
passage northwards from Tunisia.
The route around the eastern end of the Mediterranean is
used by many more Black Storks. At the Bosphorus, a total
of 6194 passed between 16 August and 25 October 1966,
and over 7400 birds were counted between 8 September and
3 October 1973 (Porter and Willis 1968, Cramp 1977:325);
the peaks came in late September. A total of 203 were seen
in a 3-h period near the Bosphorus in March 1963 (Kumer-
loeve 1966b). Parties of up to 500 birds are reported from
Israel in both spring and autumn; and, on their journey
north, 3500 were counted 26 March 1981 above the Valley
of the Moon near Eilat. Formerly a rare winter visitor to
Israel, it has been increasing in recent years, possibly be-
cause of the increased number of artificial fish-ponds. As
many as 290 Black Storks remained in the Hula and Bet
She'an Valleys in the winter of 1982-83 (Paz 1987:34). A
few birds winter as far north as Turkey (Vittery 1972).
Birds heading for Africa apparently travel down Sinai
and the Red Sea coast, for they are said to be rare in the Nile
Valley (Cramp 1977:325). Some birds have been observed
on the Red Sea, near Jiddah, and in small flocks near Aden,
but no evidence is available to show the origin of these
migrants. Wintering birds are recorded regularly in small
numbers south in Africa to the Sudan (about 100 annually),
Kenya and northern Tanzania, and these are most likely of
European origin (Backhurst et al. 1973); birds in southern
Tanzania may be of southern African origin (Britton
1980:21). Ringed birds from northeastern Europe have been
recovered in Ethiopa and Uganda (Schiiz 1940b, Bauer and
Glutz von Blotzheim 1966:422).
There is one report of a bird ringed in Germany taken as
far south as Malawi [formerly Nyasaland] (Belcher
1930:12); however, we feel this was an error, probably
referring instead to the White Stork (see also C.W. Benson
1958). And we agree with Prigogine (1976) that reports of
'large flocks' of Black Storks in tropical Africa (e.g. V.G.L.
Van Someren 1922, Verheyen 1953) almost certainly refer
to the Abdim's Stork.
Birds enter Pakistan on a broad front across northwest
Baluchistan and via the northwestern Himalayas, and a
flock of over 100 has been recorded in Gilgit in the autumn
(TJ. Roberts pers. comm.). TJ. Roberts (1969) reported
scattered sightings of birds in Sind and, in January 1969,
saw 81 birds in the Punjab of Pakistan, and they have been
noted in the Kurram valley. Certainly parties of Black
Storks winter in northern India regularly. They have been
seen at least as far south as 14-15°N in India (Madsen
72 Black Stork
1990), and once 2 birds were reported in Sri Lanka (Henry
1971:386, W.W.A. Phillips 1978:9). We saw a group of 7
birds, including 1 immature, in the Ranthambhor National
Park in March of 1989, in the company of a pair of Woolly-
necked Storks. Wardens there reported seeing flocks of up to
20 birds on passage through this tiger reserve.
Black Storks from the eastern USSR and China winter
mostly in southern China (Cheng 1976:34). In Hong Kong,
the Black Stork is an irregular and scarce winter visitor,
with a few individuals arriving for short periods, but not
every year (Chalmers 1986:41). It is an uncommon visitor
further south, in Burma, northern Thailand and Laos
(Delacour and Greenway 1940, Smythies 1953:521, B.F.
King and E.G. Dickinson 1975:51, P. Round et al. unpub.
report). In Korea, the Black Stork is a rare and local
summer visitor, occasionally wintering in the south of the
peninsula (Austin 1948:46, Gore and Won 1971:119).
Generally this stork migrates in small flocks, but indi-
vidual birds disperse nomadically and have been recorded
from as far afield as Britain, Norway, Finland, Switzerland,
Malta, Cyprus, Madeira, Taiwan and Japan. Old ringing
records .show that from a brood of 4 ringed in Denmark, 2
were recovered southwestward in the Netherlands and
northern France, and 2 southeastward in Hungary and
Romania. 2 young from an East German nest were re-
covered in France and Hungary (Cramp 1977:324). Birds
ringed in Latvia were recovered in Poland in September, in
southern Asia Minor in November, and in Egypt in March
of the following year. Birds ringed in Lithuania were re-
covered in Hungary and in the central Soviet Union. One
ringed in the Kaliningrad, USSR, region was recovered in
Peloponnesus (Greece) (G. Niethammer 1938).
Spring migrants arrive back at their nesting areas be-
tween March and May, mostly in April. In many areas they
return about 1-2 weeks later than the local White Storks (G.
Niethammer 1938); but in southwestern Poland in the
spring of 1969 Black Storks preceded the White Storks by a
few days (pers. obs. M.P.K.).
Populations and trends
Populations are small; the Black Stork is nowhere common,
and its numbers are diminishing everywhere. In the USSR,
the Red Data Book describes its status as widely distributed
but rare and numbers decreasing everywhere. The total
population in the USSR is unknown, but no more than 1QO
birds breed in the western Ukraine. In Estonia, in 1970, it
was estimated that at most 80 pairs nested, compared to 150
recorded in 1962. In Latvia, in the 1950s, there were 260
nesting pairs. About 200 pairs are said to inhabit the Kha-
barovsk region of the extreme eastern USSR (Sonobe and
Izawa 1987:43). Annually 2-3 pairs nest in Qianshan
Nature Reserve, Liaoning Province, northeastern China
(Sun 1991). The Black Stork apparently no longer occurs in
southern Korea as a breeding bird (P. Won 1973:18); how-
ever, birds were recently observed during the spring and
summer in northeastern Korea (Hamgyong-Pukto) and
could nest there (Sonobe and Izawa 1987:44).
A few pairs may still nest in western Germany, e.g. in the
Liineburger Heide (Miiller-Scheessel 1964, 1965, Makowski
1970), and 5 or possibly more pairs in Austria; in southern
Greece there are under 20 pairs. The species has disap-
peared as a breeding bird from Denmark (Hans Skov pers.
comm. 1990), where about 150 pairs bred in 1850; this
number declined to about 70 pairs in 1900, 40 in 1910, 20 in
1920, 3-5 in 1930-38, 1 (irregularly) through the early
1950s. The last successful breeding was probably in 1953.
Its disappearance in Denmark was mainly from destruction
of old forests and too much human activity near the nests;
also many ringed birds were shot during their migrations
(outside of Denmark).
In Yugoslavia, numbers have declined sharply, especially
in the north of the country, but 50-80 pairs remained in the
late 1980s along the Sava River in Croatia (M. Schneider
1988b). In Poland the situation is one of improvement with
an increase of breeding since 1936, after 50 years of decline.
In 1966, 480 occupied nests were found in Poland, and the
total population estimate rose to some 500-530 pairs; by the
mid-1980s the estimate for Poland was about 800 pairs. In
eastern Germany, 18 pairs remained after 1963, and in
Czechoslovakia there was an increase since 1940 of up to
100 pairs in 1960. There were 50-60 pairs in Hungary in
1941, and about 150 pairs in the mid-1980s. Two isolated
populations remain elsewhere in Europe: one in north-
eastern Portugal, where some 25-30 pairs still breed, and
100-150 pairs in central and southern Spain (Cramp
1977:324, Luthin 1987).
A few pairs breed in southern Africa, where the discovery
of nesting is a comparatively recent occurrence (Clarke
1904). Siegfried (1967) reports 15 pairs breeding in Zim-
babwe (formerly Southern Rhodesia), 9 in Cape Province
and 4 in Natal, South Africa, 3 in Lesotho (formerly Basuto-
land), and 1 each in Mozambique, Botswana (formerly
Bechuanaland) and Orange Free State, South Africa, a total
of 34 breeding pairs. In a later estimate, Siegfried et al.
(1976) state that there are 'probably no more than 100
breeding birds' in South Africa itself, and that this popu-
lation is stable. Transvaal Province, South Africa, supports
an additional 50-70 breeding pairs (Tarboton 1982); a few
pairs also breed in Zambia (C.W. Benson et al. 1971:46), at
least 2 pairs in Malawi (J.H. Ryder and B.A. Ryder 1978)
and probably a few pairs in the Kuiseb River Canyon,
western Namibia (Tilson and Kok 1980).
ECOLOGY
The Black Stork usually avoids human habitation and
favours well-wooded areas with shallow lakes and ponds as
well as marshlands, damp meadows, rivers, and streams. It
is very dependent on water, being more an aquatic feeder
and fish-eater than the White Stork, so much so that in
southern Africa, at least, nesting is not carried out in
drought years (Tarboton 1982). Although usually frequent-
ing freshwater areas over most of its range, non-breeding
Black Storks are often found in the estuaries of tidal rivers in
South Africa (Siegfried 1967). It may feed on drier land in
the winter, when it will take reptiles, mammals, and insects.
Unlike most stork species, the Black Stork is often found

in hilly or mountainous regions, and has nested as high as
2000-2200 m in Soviet Central Asia (Dementiev and Glad-
kov 1951:441); it is also found up to at least 2000m in
Lesotho, southern Africa (Brown et al. 1982:179) and at
about 2500 m in the western Sudan (Lynes, in Bannerman
1957:20).
Feeding is by deliberate stalking while Walking Slowly in
shallow water, either singly or in small groups. Food loca-
tion is mainly visual, and the bird Probes vertically and
grabs prey in the tip of the bill with a quick forward lunge of
the head. Breeding adults in Poland may fly up to 10km
from the nest site to gather food (Cramp 1966).
On rare occasions Black Storks have been observed forag-
ing with both wings raised in an open canopy (England
1974, Breife 1982). This has been compared with the Black
Heron (Egretta ardesiaca), which shades the water with open
wings and darts its head forward to catch prey, but although
the wings are bent and pushed forward of the normal flying
position the Black Stork at no time forms the closed canopy
characteristic of the Black Heron.
Fish makes up the main food of this bird; species captured
include loaches (Misgurnus and Cobitis], pike (Esox lucius),
burbot (Lota lota), rudd (Scardinius erythrophthalmus], roach
(Rutilus mtilm\ perch (Perca fluviatilis], eel (Anguilla anguilla)
and sticklebacks (Gasterosteus sp.). They have been seen to
catch and swallow fish up to 30 cm long (Meinertzhagen
1954:390). And they also sometimes eat insects, frogs, snakes,
crustaceans, small mammals, lizards and nestling passerine
birds (Cramp 1966, 1977:326, Bauer and Glutz von Blotz-
heim 1966:427, Dementiev and Gladkov 1951:444, Witherby
et al. 1939:117, Schroder and Burmeister 1974:26). In Africa,
the Black Stork is less dependent on insects than White or
Abdim's Storks and does not commonly follow locust swarms
or outbreaks of army worm (Spodoptera sp.). In Pakistan,
TJ. Roberts (1969) reported them eating snails (Vivapara
bengalensis and Lymnaea acuminata).
This species is much shier and more solitary than the
White Stork. Birds generally nest and feed in secluded
places, rarely visited by man. Dementiev and Gladkov
(1951:444) describe it as 'exceedingly circumspect', and
Siewert (1955:48), who studied the species intensively at the
nest in northeastern Poland, referred to it as a Kulturfluchter
(a fleer from civilization). However, in the Transcaucasus
they are known to feed in cultivated areas, and here they
also nest close to human habitation, occasionally even in
Azerbaijani hamlets (Dementiev and Gladkov 1951:441).
While occasionally seen in flocks of up to 100, it is more
usually observed singly or in small parties of 3 or 4 birds.
Predation by eagles (Aquila sp.), especially during mi-
gration, has been reported (Ritzel 1980, Madsen 1990).
BREEDING
Nests are often used year after year and the pair-bond may
be of long duration; in Europe it is rare for a given nest to be
used for more than 10 years (Schroder and Burmeister
1974:16). Pairs often return to their traditional nest site
together, and it may be that pairs migrate and pass the
winter together.
Black Stork 73
Usually the Black Stork nests solitarily, and where it is
numerous, nests are usually at least 1 km apart. In the
central part of Transvaal Province, South Africa, the closest
nests were 6-20 km apart; in Kruger National Park, eastern
Transvaal, some nests were 2.2-4.7 km apart (Tarboton
1982). Nesting in Europe is usually in old, large trees (e.g.
beech, oak, pine) in deep forests, but may be on a wooded
river bank or, especially in Asia or Africa, on a cliff. Terri-
tory near the nest and surrounding area is defended against
conspecifics. Contrary to other records, two occupied nests
in the same tree have been reported in the Lenkoran Low-
lands, Azerbaijan, USSR (Dementiev and Gladkov
1951:441).
In Europe, nests are most often 4-25 m high in trees—a
wide variety of both evergreen and deciduous; in many parts
of Asia and Africa, nests may be in niches on cliffs, caves,
large boulders, or under overhanging ledges. Usually nests
are well away from human habitation. Some are high up in
trees, which can be seen from long distances, whilst others
are well hidden. The trees selected are usually very large
and the nest is often on a large horizontal branch, about
two-thirds of the way up the tree. Dementiev and Gladkov
(1951:442) found that near Lenkoran, Azerbaijan, the storks
nest on the outskirts of colonies with herons and cormor-
ants, though this must be considered unusual. In Transvaal,
South Africa, they sometimes nest in colonies of the Bald
Ibis or near Cape Vultures (Gyps coprotheres), Black Eagles
(Aquila verreauxi], Lanner Falcons (Falco biarmicus), or Pere-
grines (Falco peregrinus] (Tarboton 1982). In exceptional
cases, nests have even been built on the ground (Bauer and
Glutz von Blotzheim 1966:423).
The nest is a large structure of sticks and twigs, and
sometimes quite large branches. Old nests that are used
again and again in traditional nesting sites are repaired and
characteristically reinforced with earth and grass, and lined
with moss, leaves, grass, paper, animal-fur and rags. Some-
times old raptor nests are taken over. In southern Africa,
they occasionally nest atop Hamerkop (Scopus umbretta)
nests, or those of the Black Eagle (Maclean 1985:66). Old
Black Stork nests can be 1-2 m in diameter, but are con-
siderably smaller if newly constructed (Dementiev and
Gladkov 1951:442; Brown et al. 1982:179). In general, they
are somewhat smaller and shallower than nests of the White
Stork (Bauer and Glutz von Blotzheim 1966:423).
Unless the pair arrives together, the male usually returns
first to the nest site. Threat displays and fighting are less
common than in the White Stork. Birds often spread their
long, white undertail-coverts and press them and the black
tail (not spread, but compressed into a c. 12 cm wide panel)
downward to about the vertical during displays; in hostile
encounters, the luxuriant feathers of the lower foreneck are
fully erected, giving the bird a larger, fiercer appearance
(Siewert 1932a, Bauer and Glutz von Blotzheim 1966:426,
pers. obs. M.P.K.).
A variation of the Up-Down display is given towards
intruders at the nest, in which the bill is lifted to the vertical
and clattered (Stoll 1934). The Up-Down given as a greet-
ing between mates is similar but less vigorous and most
often without the vertical bill-movements and clattering.
74 Black Stork
According to several observers, bill-clattering is only rarely
recorded in Black Storks in Europe (O. Heinroth and M.
Heinroth 1924-31:141, Stoll 1934, Siewert 1932b:35,
1955:70, Rosenberg 1941), but we have seen it during the
Up—Down display between mates in South African birds
(Kahl 1972c). Up-Downs are accompanied by a series of
weak, melodious bisyllabic hissing whistles, synchronized
with the tossing movements of the head and sinuous move-
ments of the neck.
Although the Up-Down display of the Black Stork is very
different, some other mating displays are similar to those of
the White Stork, including Head-Shaking Crouch, Copula-
tion Clattering and Display Preening. Before copulation, the
female often bends forward, with her body about horizontal,
and presses up against the breast of the male; he sometimes
hooks his bill over her back before stepping on (pers. obs.
M.P.K.). After copulation, mutual Up-Down displays and
allopreening are often performed. Before egg-laying the pair
spend much time on the nest together, with bills resting on
or buried in the luxuriant neck feathers.
Nest-building is done by both sexes. This usually involves
the repair of an old platform, from previous seasons, and the
adding of soft material to the lining.
As long as 2-3 weeks may elapse between the arrival of a
pair at the nest-site and the laying of eggs (G. Niethammer
1938:303). Clutches vary between 2 and 5 eggs, rarely 6. For
82 clutches in Poland, the mean was 3.2 eggs/clutch (Mru-
gasiewicz in Cramp 1977:327); in southern Africa, 90
clutches averaged 2.8 eggs/clutch (Brown et al. 1982:179)
and another 19 clutches averaged 3.4 eggs/clutch (Tarboton
1982). The eggs are smaller than in the White Stork, and the
shell is white with a greyish hue; they become soiled as
incubation progresses. Eggs are laid at 2-day intervals, and
incubation, by both sexes, commences after the first or
second egg is laid. The incubation period is variously given,
as 30, 35-36, 32-38 or 35-46 days (Dementiev and Gladkov
1951:443; Bauer and Glutz von Blotzheim 1966:424, Cramp
1977:327); the lower figures are probably the more accurate.
Hatching is asynchronous. At least one adult remains on
the nest, to guard eggs or young, until the nestlings are 2-
3 weeks old (Ruthke 1957, Cramp 1977:326,328). Both
parents feed the young by regurgitation onto the floor of the
nest. At Oka sanctuary, USSR, young were fed mainly on
small (9-25 cm) fish and a few frogs; each parent fed 3-8
times daily and the daily consumption of each nestling was
400-500 g (Cramp 1977:326). Begging young utter monoto-
nous braying calls similar to other stork species, and clatter
their bills when aggressively excited (Stoll 1934). Nest pre-
dation is apparently uncommon; pine martens (Martes
martes) occasionally take eggs or young (Ruthke 1957).
Fledging takes about 60-71 days, and after that the young
join the adults at their feeding grounds. Young continue to
return to the nest for another 2 weeks, to be fed and to roost
at night (Ruthke 1957, Bauer and Glutz von Blotzheim
1966:424). Nesting success varies from 31 to 92% from year
to year, with about 3.4 young/nest in successful pairs. One
pair in Czechoslovakia successfully reared 5 young (Stoll-
mann 1961). Birds apparently stay together in family
groups until migration (Dementiev and Gladkov 1951:443;
Cramp 1977:328). G. Niethammer (1938:302) considered
that sexual maturity is reached at the age of 3 years, because
a bird ringed as a nestling bred at that age.
In northern South Africa, Tarboton (1982) found that
pairs failed to breed in 38% of the years, usually when
rainfall was below average.
TAXONOMY
For many years most classifications listed the Black Stork
and the White Stork (including the Oriental White Stork) as
the only members of the genus Ciconia. We, however, in-
clude the Abdim's, Woollynecked, Storm's and Maguari
Storks in the genus Ciconia (Delacour and Mayr 1945, Kahl
1972e). Detailed study of the courtship behaviour of this
group (Kahl 1972c) suggests that the Black and White
Storks may be the least closely related species in the genus,
with the others, formerly placed in separate genera, in
intermediate positions between them.
In spite of its extensive range, no subspecies of Ciconia
nigra have been described. The birds in southern Africa are
probably a comparatively recent offshoot from the Palaearc-
tic population; they seem to be reasonably well isolated,
reproductively, but have not yet developed any noticeable
differences from the parent-group (Moreau 1966b:122,
Snow 1978:30).
CONSERVATION
The Black Stork is a rare and threatened species throughout
most of its range. It is undergoing a long-term population
decline in western Europe, and habitat changes are proceed-
ing at a rapid rate in much of eastern Europe and Asia. It
has already been extirpated from Scandinavia and nearly so
from France, West Germany and Greece (Luthin 1987). In
eastern Denmark, for example, the population was esti-
mated to be 150 pairs in 1850, but had decreased to half of
that in 1900, and to 20 pairs in 1920 (E.V. Rasmussen
1979); the last stork nested there in 1953. Although Black
Storks are seen each year, they no longer breed in Denmark.
This has been the tale throughout western Europe.
The main threat to this species is the destruction of
forests, particularly large trees on which nesting tradition-
ally has occurred. In Russia and other parts of eastern
Europe, the rapid development of industry and farming has
reduced areas of breeding. Heavy predation by man in
southern Europe and tropical Asia, during migration, have
reduced the population considerably. In addition, habitat
changes, particularly the loss of wetlands and the use of
pesticides, in wintering grounds in Africa will undoubtedly
adversely affect the Black Stork as they have the White
Stork. According to Sun (1991) nestlings in northeastern
China are often stolen for sale to zoos.
The Black Stork is a wilder, shier creature than the White
Stork, but without full protection on its breeding grounds,
and during its extensive dispersant and migratory journeys
it will inevitably fall rapidly in numbers throughout its
range.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 290.
 Abdim's Stork
Ciconia abdimii Lichtenstein

Ciconia abdimii Lichtenstein, 1823, Verzeichniss Doubletten Zool. Mus. Berlin, p. 76: Dongola, Sudan

Other names:
Whitebellied Stork, Smaller Locustbird (English); Abdimstorch, Regenstorch (German); Cigogne
d'Abdim (French); Abdim-ooievaar, Regenooievaar (Dutch); Kleinswartooievaar, Blouwangooie-
vaar, Swart Sprinkaanvoel (Afrikaans); Sinbilah (Arabic); Shamuwa, Shamwona (Hausa); Eviyoyo
(Kakamega); Omenena (Luo); Shuramurove (Shona); Mokoroane, Lekololoane, Roba-re-bese
(South Sotho); Lekololwane, Mokotatsie (Tswana); Endongondongo (Kwangali)

IDENTIFICATION
Abdim's Stork is a comparatively small member of the 'typical' stork group, standing 68-81 cm tall. It
is predominantly blackish, with a white rump and belly.
The adult is a predominantly black bird, with the dark head, neck, upper breast, and back glossed
with purple and green; the tail and wings are also black. Its lower back, including the uppertail-coverts
and underparts, from the lower breast to the belly are white. At rest, a narrow line of white breast
feathers shows between the bend of the wing and the throat.
The iris is greyish brown, and the red-tipped bill is otherwise a pale horny jade-green. The bare skin
of the face is lead-blue, with a red patch over and in front of the eye and under the bill. The legs and feet
are dull olive, with a reddish 'garter' at the tibio-tarsal joint and on the toes. The male is slightly larger
than the female, and the male's bill is deeper and thicker at the base (Brown et al. 1982:180).
At the commencement of breeding, most of the bare facial skin becomes bright French blue and the
rest of the soft parts assume a richer colour. The lores, eye-ring, chin and inside the ears are vermilion-
red, the forehead a lighter flesh-colour, and the remainder of the naked face bluish-purple. Breeding
birds have greenish grey legs, with pinkish red intertarsal joints and toes (Andersson 1872:280, A.L.
Butler 1905, Kahl 1972c).
Two captive young took 32-36 h from pipping to hatching. Newly hatched young weighed about 37 g;
they are thinly covered with dark down, over pale blue-grey skin, and have black foreheads. By 10 days,
nestlings are covered with soft light greyish down, the bill is black with a white tip, and the legs are
pink. Half-grown nestlings are covered with whitish down, except for the front of the crown where the
down is black; this head and neck pattern—as shown in the photo in Bigalke (1948) —betrays a close
relationship to the Woollynecked Stork, which retains this pattern in the adult. By 4 weeks of age, the
young are fully feathered and resemble the adults, although substantially duller (North 1940, Bigalke
1948, Farnell and Shannon 1987).
Immature birds are patterned like adults but are much duller and browner, with less gloss, and less
colourful soft parts. There is some down on the bare skin of the throat, and the bill is a dull reddish
(Mackworth-Praed and Grant 1970:54).
The bisyllabic whistles are similar to those uttered by the Black Stork. Calls given at a roost in
Zimbabwe have been described (Brooke 1969) as 'solitary whispered whistles which carry some
25 yards'. 'For a stork its cry is most trivial —a high-pitched "peep . . . peep . . . peep . . .", worthy of a
little more than a barn-door chicken' (Lynes 1925a:550).
Flapping flight is used for short-distance travel, and soaring/gliding flight for longer distances,
especially on migration. Our observations at Lake Shala, Ethiopia, demonstrated the pattern that is

general for most storks. Abdim's Storks left the nesting
island to fly, by flapping flight, to the mainland. On the way
back, they were first able to gain altitude in the thermals
over the land, and could soar and glide back to the island.
However, if the wind was high and soaring not possible,
they returned by flapping flight low over the water (Kahl
197Id). We have seen similar situations in several species of
storks. Abdim's Storks engage in incredible acrobatics when
descending from high altitudes (Mackworth-Praed and
Grant 1962:81). In normal flapping flight, the flap-rate was
189/min(n = 28)(Kahll971b).
Abdim's Stork could be confused with the more svelte
Black Stork, but it is a considerably smaller and more
'dumpy' bird, shows a white rump and lower-back in flight,
and differs in its soft-part colours. It is gregarious at most
times, unlike Black Storks, which are seldom seen in flocks.
The White Stork differs, of course, in being predominantly
white. And the Woollynecked Stork is somewhat taller and
longer necked, with a whitish head and neck. Other African
storks are larger and distinctive in their own ways. Sympa-
tric herons are thinner and differently coloured, have a
different 'jizz' (Gestalt^ manner), and are found in more
aquatic habitats.
DISTRIBUTION AND POPULATION
It nests in the northern semi-arid tropics from Senegal,
northern Nigeria and Chad, eastward through Sudan, Ethi-
opia, northern Somalia, southward to western Kenya and
Uganda; it also nests, in small numbers, in Yemen, south-
western Arabia (Dekeyser 1952, Meinertzhagen 1954:391,
K.D. Smith 1955a, Montfort 1965, Moreau 1966b:247,
Urban 1970, Kahl 1971d, 1979b, Gallagher 1986). In West
Africa, its breeding is largely confined between 17° and 8°N
(Bannerman 1953:172); in East Africa it breeds, in small
numbers, south to the equator.
Abdim's Stork is an intra-African, trans-equatorial mi-
grant, breeding north of the equator and wintering mainly
Abdim's Stork 77
to the south. Its migratory routes and times of movement
are well recorded. Movements seem largely timed to follow
the rains, and their associated insect abundance (Andersson
1872:280), back and forth from northern to southern hemi-
sphere. This stork rarely spends much time in dry con-
ditions (Moreau 1966b:246, Brown et al. 1982:180).
Birds in a region concentrate in large numbers in prep-
aration for their southward migration; in Sudan this occurs
in August and September (Luthin 1984a). Birds from West
Africa move eastward and then southward by September-
November; they take a more easterly route than in the
spring and probably fly at great heights over the Congo
forest (Chapin 1932a:457, Bannerman 1953:173). From
eastern Africa, they fly south often in huge flocks, becoming
a passage migrant in September-December in Uganda,
western Kenya and western Tanzania. The vast majority of
these birds spend the winter (Dec-Feb) further to the south
(Britton 1980:20). Flocks arrive in South Africa in October,
departing northwards towards the end of March; they are
most frequent in the drier inland areas (i.e. Orange Free
State, Transvaal), and also in Botswana and northern
Namibia (Clancey 1965:229, McLachlan and Liversidge
1970:52). Up to 800 birds have been reported from Zambia
in February and early March (C.W. Benson et al. 1971:46).
They pass northward through Uganda, western Kenya and
western Tanzania in late February to May. Abdim's Stork is
said to migrate at night (G. Archer and E.M. Godman
1937:68), but we doubt if this is common.
A migrant Abdim's Stork was captured in Zimbabwe
carrying an arrowhead imbedded in its chest. The arrow-
head, which was surgically removed, proved to be of a type
originating in the eastern portion of West Africa. Later the
bird was released in apparent good health (Condy 1966).
Some birds, mostly subadults and immatures, wander
irregularly eastward to Oman, eastern Arabia; this move-
ment may be increasing owing to expansion of cultivation
and irrigation (Gallagher 1986). It has been recorded from
southern Spain as a vagrant (Heuglin 1869-73:1107). There
appears to be a small, sedentary (i.e. non-migratory) popu-
lation in eastern Eritrea, Ethiopia (K.D. Smith 1955a, Snow
1978:31); they breed during the January-March winter
rainy season, when most other Abdim's Storks are in the
southern hemisphere.
The species is common to abundant throughout its range,
at times gathering during migration in groups of thousands
at appropriate feeding sites. Breeding colonies are generally
small, a few dozen birds at the most, but the breeding range
is extensive and total numbers are at least in the tens or
hundreds of thousands. In January 1987, Donald A. Turner
(pers. comm. 1990) observed over 40000 Abdim's Storks, in
company with over 100000 White Storks, feeding on an
army-worm infestation, approximately 50 km west of
Arusha, Tanzania. Overall, its populations are considered
to be stable (Brown et al. 1982:180).
ECOLOGY
This stork is a terrestrial species, seldom feeding in water,
and most commonly found in grasslands, pastures and culti-
78 Abdim's Stork
vated fields. It is primarily insectivorous, and its migration
patterns can be altered by the local appearance and distri-
bution oflocust swarms, as storks move to take advantage of
this food resource. According to Andersson (1872:280), 'The
more plentiful the rain, the more abundant the birds, the
cause being simply the greater abundance of food.' Burnt
ground, where new grass grows after rain, also supports
high densities of readily available insects. Abdim's Storks
are also attracted to grass fires, where they patrol near the
flames.
Foraging is mainly diurnal and food-location primarily
visual. Abdim's Storks generally feed in loose aggregations.
At fires and on burnt ground, they often are in the company
of Marabou and White Storks, kites, rollers and other birds.
At other times, small parties of this stork will Walk Slowly
across dry fields with necks stretched forward and heads
erect, looking for their prey. At locust swarms, flocks of
thousands can be seen. Under these conditions, a stork
Walks Quickly, picking up insects until satiated. As Condy
(1965) observed, single birds or pairs feed in a more leisure-
ly manner, whereas birds in larger groups seem stimulated
to forage at a faster rate. Is this because of social stimulation
or just better foraging conditions (e.g. more food)? In Zim-
babwe, a 'fair-sized flock' of Abdim's Storks, along with 4
Pied Crows (Corvus albus), reportedly cleared a 4-acre (1.6-
ha) field that was heavily infested with army-worm cater-
pillars in 3 days (Hamling 1953).
The Abdim's distinctive diet is made up primarily of
insects, and it is best known for its exploitation of plague-
insect swarms. It has been known to take frogs, lizards,
other small reptiles, small fish, water-rats, young birds,
millipedes, scorpions, snails and crabs. Its insect prey in-
cludes beetles, grasshoppers, crickets and army worms (Spo-
doptera sp.), caterpillars, and especially locusts such as the
red locust (Nomadacris septemfasciata) (Andersson 1872:280,
Heuglin 1869-73:1107, Ghapin 1932a:458, Meinertzhagen
1954:391, Dean 1964, Brown et al. 1982:180).
This species has been reported (Friedmann 1930:30,
Curry-Lindahl 1961) foraging near large mammal car-
casses; it was not determined whether they were actually
eating carrion or just the insects attracted to it (most likely
the latter). When not feeding, flocks often gather near pools
or water-holes. Sometimes the birds sun themselves with
wings extended widely (North 1940). Soaring is common at
midday near feeding areas and may aid thermoregulation
during hot weather (Brown et al. 1982:180).
There appears to be little aggressive action between these
birds, but occasionally they will use Forward Displays or an
Upright Display, with bill held high and body feathers
compressed and perhaps with the neck ruff intermittently
erected for a few seconds (Kahl 1972c). Often Abdim's
Storks associate in feeding groups with White and Woolly-
necked Storks. And sometimes they are followed, or even
ridden upon, by Carmine Bee-eaters (Merops nubicus) (Rei-
chenow 1900-01:345, Cunningham-Van Someren 1970).
This species is often extremely tame around nests, yet
much less so when feeding away from the nesting area (A.L.
Butler 1905). At the colony at Lake Shala, Ethiopia, one
could approach almost to within touching distance of birds
on nests before they flew off; even then they moved only a
few metres (Kahl 197Id). Local people generally do not
bother the Abdim's. However, they are reportedly eaten by
the Ennedi Nomads, in Chad (Salvan 1967); being insecti-
vorous, they are presumably rather tasty.
BREEDING
Abdim's Stork is a colonial breeder, unlike other 'typical'
storks (genus Ciconia). Breeding groups are usually small,
consisting of a few to a few dozen pairs; as many as 30 nests
have been reported from one mimosa tree in the Sudan
(Brehm 1854). They sometimes nest in mixed colonies, with
Pink-backed Pelicans (Pelecanus rufescens), Marabou Storks,
Cattle Egrets (Bubulcus ibis), and Sacred Ibises nesting
alongside each other (Lynes 1925a:550, Kasoma and
Pomeroy 1987). They have been found nesting as high as
2100m in northwestern Ethiopia (Cheesman and Sclater
1935).
Breeding is during the rainy season north of the equator,
coinciding the time when insect populations are at their
most prolific. In Ethiopia we found that the actual initiation
of nesting coincided with the first heavy rains of the season,
which produced a rush of chironomid flies and probably the
insect prey of the storks as well. Heavy rain a week and a
half later was correlated with a marked increase in courtship
and nest building activity (Kahl 197Id). Thus it seems to us
that rains or the food supplies they produce are the proximal
trigger of courtship and nesting. An exception to the 'spring'
nesting season is in equatorial East Africa and coastal Eri-
trea, where heavy rainfall occurs at other times of year
(K.D. Smith 1955a, FJ.Jackson 1938:73).
Abdim's Storks build their nests in trees, such as baobab,
acacia and palm, as well as on rooftops of African huts;
sometimes on masses of rock, such as on the columnar
basalt formations of Lake Shala, Ethiopia (Kahl 197Id). In
northwestern Kenya, Blencowe (1962) found them nesting
on cliff ledges. Particularly in West Africa, nests are often
built in trees in villages or even on the roofs of huts, and
tribesmen welcome their annual return as harbingers of the
rainy season. At Lake Shala, Ethiopia, we found that the
nests were close together, some actually touching, although
most were separated by 30 cm or more. They are made of
sticks and lined with softer vegetation, and are 1.0-1.5 m in
diameter and 20-30 cm thick (Kahl 1971d, Brown et al.
1982:181).
Nesting begins with the male establishing a territory
either at an old nest or at a new site. The frequency of
ritualistic displays is greater than in most other members of
this genus, probably because it is the only member to nest
colonially. During the selection of nest sites Aerial Clatter-
ing is very commonly used. The attacker flies after an
opponent, attempting to maintain a position above and
behind. As the attacker nears its opponent, it lifts its head
slightly above the horizontal and clatters the bill loudly (up
to 25 times).
An unusual display by males before and during pairing is
termed Mock Fighting (Kahl 1972c). This rather bizarre
display seems sometimes to be triggered by an insect flying

near the bird; at other times it appears to be spontaneous. It
erects the neck, head and breast feathers and makes a series
of wild lunges in the air, snapping and rattling the bill. It
leans from side to side as though buffeted by wind, often
losing its balance, flaps its wings to retain balance, and bites
vigourously at nearby objects or nesting material. Some-
times this display is followed by the bird diving steeply from
its perch and flying around erratically in a darting, bat-like
flight, sometimes with the feet thrust forward in the manner
of a bustard on take-off. In a letter to Margaret Nice,
Konrad Lorenz describes this display precisely (Nice
1943:68) in captive Abdim's and Black Storks (it has not, to
our knowledge, been reported in wild Black Storks): 'When
[they] are let out into a larger enclosure . . . they begin a
curious dance, evading by "hooks" and sudden ducking the
non-existent attacks of an imaginary eagle, even thrusting at
him vertically upward with their bills.' Cathy King (pers.
comm. 1990) has observed a similar display in captive
Maguari Storks.
When an unmated female approaches a male at a poten-
tial nest site, the Head-Shaking Crouch is frequently per-
formed. The male utters a series of bisyllabic whistles and
inclines its body forward, with head lowered and bill down.
The wings are held slightly away from the body and lifted.
Back and neck feathers are erected and the tail cocked. As
weight is shifted from foot to foot, the head is shaken from
side to side, and then the bird lowers its body to lie flat on
the nest. Sometimes the approach of the female after such a
display will result in her being driven off. Only occasionally
is this display performed by a newly mated female as her
mate approaches the nest, and after pair-bonding is com-
plete is it no longer carried out.
The Up-Down display of this stork varies strikingly from
the well-known display of the White Stork. Throughout the
display, the wings are spread and held at an angle of 90°.
The bill is first pointed downward, with the neck arched; a
series of melodious whistles are given. Then the head is
raised until the bill is pointing upward, and the mandibles
are clattered rapidly a number of times. The tail is alter-
nately lifted and depressed, with head movements. To our
knowledge, the Nest-Covering display has not been de-
scribed for this species.
The very active displays of Abdim's Stork at the time of
pair-formation are followed by the completion of the nest or
addition to a used nest from the previous year. The male
brings most of the material, which the female uses to build
the structure. Completed clutches are 1-5 eggs, usually 2 or
3; 16 clutches averaged 2.2 eggs (Brown et al. 1982:181).
They are laid at 2 or 3 day intervals. Eggs are chalky cream-
white and soon become soiled.
Data are not available for incubation periods in the wild.
Abdim's Stork 79
The incubation period of a captive pair was 30-31 days, and
began with the laying of the second egg (Bigalke 1948);
artificial incubation of 2 eggs from another captive pair took
28-30 days to hatching (Farnell and Shannon 1987). Both
parents incubate and continue to accompany nest exchange
with Up-Down displays. The only egg mortality noted at
Lake Shala was caused by intraspecific fighting at the nests
(Kahl 197 Id).
Adults feed young by regurgitating onto the nest. Parents
often shade nestlings from the hot sun, with outstretched
wings. Predation is avoided mainly by one parent being
present while the nestlings are small. Eggs and chicks are
lost through disputes of neighbouring birds, and attempts to
take over nests or nesting material. The chicks develop
slowly, being able to sit by the sixth day, and standing at
2 weeks. They are fully feathered by 4 weeks of age (Farnell
and Shannon 1987). The fledging period is probably 50-
60 days, and the entire breeding cycle, from arrival of adults
to the flight of the young, takes about 90-100 days (Brown
et al. 1982:181). According to Bannerman (1953:173) 3-4
young, rarely 5, are reared in a season.
TAXONOMY
Once placed in the monotypic genus Sphenorhynchus, and
before that in Abdimia, the Abdim's Stork is now grouped
with the other 'typical storks' in the genus Ciconia (Delacour
and Mayr 1945, Meinertzhagen 1954:391, Kahl 1972e). It
was named after Abdim Bey, the Governor of Dongola,
Sudan, in 1823 (Kahl and Schiiz 1972).
CONSERVATION
This locally abundant stork is fortunate in that native Afri-
cans hold local individuals in high regard, realizing that
their arrival signals the beginning of the rainy season (G.
Archer and E.M. Goodman 1937:68). Of its significance to
the local Africans, Lynes (1925a:550) states: 'If Abdim does
not bring baby boys, at least it brings the precious rains'. It
is also recognized as an aid to pest control, owing to its large
intake of insects. Threats include habitat loss and pesticides.
In the Sudan, it has been suggested that pesticides are
adversely affecting young, a suggestion that needs addi-
tional study (Luthin 1984a). Efforts to control locust pla-
gues may reduce the food available to these storks.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 291.
 Woollynecked
Stork
Ciconia episcopus (Boddaert)

Subspecies:

Ciconia episcopus episcopus (Boddaert)

Ardea Episcopus Boddaert, 1783, Table des Planches Enlumineez d'Histoire Naturelle, p. 54:
Coromandel Coast, SE India
Ciconia episcopus microscelis Gray
Ciconia microscelis G.R. Gray, 1848, Genera Birds, 3, p. 561, col. pi. 151: no locality given (Africa)

Other names:

Whitenecked Stork, Whiteheaded Stork, Bishop Stork, Parson-bird, Beef-steak bird [India]
(English); Cigogne episcopate, Cigogne a cou laineux, Gigogne eveque (French); Wollhalsstorch
(German); Witnekooievaar, Bisschopooievaar (Dutch); Wolnek-ooievaar (Afrikaans);
isiThandamanzi (Zulu); Laglag (Hindi); Kali-tul (Gujarati); Manik-jor (Bengali); Vannati-narai,
Pathiri-kokku (Tamil); Karim-kokku (Malayalam); Padili-kokka (Sinhalese); Ghi-gyin-sut
(Burmese); Sandang-lawe (Javanese)

IDENTIFICATION
The Woollynecked Stork, a medium sized stork, standing 75-92 cm tall, is mostly black, with a
distinctive downy white neck, which gives it its name, and a black skull-cap. In the adult, the hindhead
and neck are white, with a black 'parson's cap'. The lower belly and undertail-coverts are white and
contrast strongly with the rest of the glossy black plumage. There is a conspicuous ruff of fluffy
elongated (c. 150cm) black feathers, with paler tan or iridescent greyish purple tips, on the lower
foreneck, which can be erected during displays (Harvey 1972, pers. obs. M.P.K.). The black portions of
the plumage are tinged with glossy greenish blue and purple, brightest coppery purple on the breast,
sides of breast, and lesser secondary wing-coverts. The deeply forked tail is black, but is often obscured
by the protruding, white, stiffened undertail-coverts, giving the impression that they are the true tail
(Priest 1933:95, FJ. Jackson 1938:71, Henry 1971). [It has sometimes been erroneously stated (e.g.
Blanford 1898:371, E.C.S. Baker 1929:324, H.C. Robinson and F.N. Chasen 1936) that the upper tail-
coverts—rather than the tail itself—are forked. This has led to some confusion.]
The legs and feet are dull red, darker on the tarsal joints and toes, the naked skin of the forehead,
orbital area, cheeks and throat is dull bluish or lead-grey; the iris is dark red or brownish scarlet
(Chapin 1932a:458, S. Ali 1953:404, Henry 1971:381). The bill is black, showing red along the culmen,
at the commisure, and often toward the tip.
In some specimens, the bill is largely dark red, with only the basal third black. This is placed by some
as a separate race, neglecta (J.H. Riley 1925:28, Chasen 1935), but we have not recognized it here. The
African subspecies averages slightly smaller and has the edges of the black cap diffused, forming a
rather jagged border of mixed black and white feathering, compared to the sharply delineated border
between black and white on the Asian subspecies. The sexes are alike, with the male somewhat larger.
82 Woollynecked Stork
The breeding plumage is cleaner looking and glossy areas
of black feathering are brighter. The iris is deep crimson or
wine-red with posterior border of the sclerotic membrane
bright yellow (E.C.S. Baker 1929:324, Bannerman 1930:98,
Chapin 1932a:458, J.H. Riley 1938:36); this conspicuous
yellow skin enlarges slightly during some courtship displays
(pers. obs. M.P.K.). At close range and when the wings are
opened, as in Up-Down display, a narrow band of brilliant
neon orange-red unfeathered skin is visible along the under-
side of the forearm (radius/ulna), it is almost glowing, like a
red-gold jewel (Legge 1880:1119, Meyer and Wiglesworth
1898:809, Bannerman 1930:98, Hachisuka 1931-32:349,
pers. obs. M.P.K.).
Small nestlings have pale greyish down over plumbeous-
coloured skin, buffy down on the neck, and a black crown.
Later the neck is thickly covered with a white down, and the
sides of the abdomen and undertail-coverts are white, while
the upper parts, lower neck and breast are a dark sooty grey
down with minute grey tips. The head, face and throat are
feathered forward to the base of the bill. The bill is brownish
black, tipped with reddish orange or yellow, and the iris is
dull brown or hazel to dull green. Legs are fleshy plum-
beous, speckled with dark brown (Whistler 1918b, Chapin
1932a:458, J.A. Scott 1975, Anthony 1977, pers. obs.
M.P.K.).
The immature bird's patterning is similar to the adult's,
although the forehead is feathered and sometimes streaked
with black and white (Clancey 1964a:45, Mackworth-Praed
and Grant 1970:54, J.A. Scott 1975). The dark portions of
the body and wings are dull brown, rather than black, and
lack the iridescence of adults. The feathers of the neck are
longer and more fluffy (Kuroda 1936:540, S. Ali and S.D.
Ripley 1968:98).
Bysyllabic whistling calls, very similar to those of the
Abdim's and Black Storks, are given during Up-Down
greetings at the nest (Kahl 1972c). It has been reported to
utter a fierce hissing when attacked by a trained falcon
(Dharmakumarsinhji 1955:91). Bill-clattering is heard, in-
frequently, during courtship displays (Kahl 1972c) and in
threat to raptors near the nest (J.A. Scott 1975). We are
sceptical of reports of 'harsh and raucous' calls (e.g.
Mackworth-Praed and Grant 1962:80, Maclean 1985:67).
When Begging, young are said to utter a call 'like a file
being used on a metal pipe' (Brown et al. 1982:182).

The Woollynecked Stork has to run for a distance along
the ground before taking off (Rabor 1977:27). In normal
flapping flight, the flapping rate averages 160/min (n = 10)
(Kahl 1971b). They often soar on thermals. 'The powers of
flight of this stork are remarkable . . . [it performs] compli-
cated evolutions, rolling and tumbling in the air and diving
at a steep angle, when the wind passing through its quills
makes a loud noise' (Bannerman 1953:171). Birds have
been observed diving from the nest and flying away in a
darting, bat-like flight (pers. obs. M.P.K.).
Woollynecked Storks often associate with Black Storks,
particularly during their extensive migrations in northern
India and Africa, but the former is easily distinguished by
its white neck. In limited areas of peninsular Thailand and
Malaysia, and southern Sumatra, the Woollynecked Stork
may occur with the very similar Storm's Stork; the former is
distinguished by the more extensive white on neck, darker
bill and facial skin; difficult to confuse with any other
species.
DISTRIBUTION AND POPULATION
This stork has a wide distribution in tropical Africa, Asia
and associated islands. It ranges from Senegal and Ethiopia
to Angola and eastern South Africa; then from Pakistan,
India and Sri Lanka to Vietnam, Thailand, Philippines, the
Sunda Islands, Celebes (but not Borneo).
The episcopus subspecies is found in appropriate habitat
throughout most of the Indian subcontinent, though Tom
Roberts (pers. comm.) considers it now extinct in Pakistan.
Formerly (Delacour and Jabouille, 1931:86) it was common
in Kampuchea, southern Laos and Vietnam. It is still wide-
spread in the southern part of Vietnam (Vo Quy, 1975) and
several were seen in central Kampuchea, as recently as 1968
(pers. obs. M.P.K.). It occurs in the Malay Peninsula, at
least as far south as southern Thailand, but there are no
breeding records for Malaysia (Medway and Wells
1976:94). No recent breeding has been recorded from Thai-
land, Burma or Bangladesh (Bain and Humphrey 1980,
M.A.R. Khan 1984, Luthin 1987). It may be quite wide-
spread in portions of Sumatra and Sulawesi (Holmes 1977,
Luthin 1987) and was, at one time, widespread in the
Philippines (duPont 1971:23) but its present status there is
not known. It also occurs in Java, Bali, Sumbawa, Flores
and Lombok.
The microscelis subspecies is confined to Africa where it is
found sparingly from the Gambia River and the Sudan
south to northeastern South Africa, Angola, and northern
Namibia and Botswana; it is less common at the southern
edge of its range. In East Africa, it is local and irregular
(Short et al. 1990:73); we have found it primarily on the
coast in Kenya, and it is not uncommon in coastal Tanza-
nia, including Pemba Island, and in the western parts of
Uganda.
It seems certain that this species migrates, but its move-
ments are not well documented or understood. When dis-
persing or migrating it will sometimes gather together in
sizeable flocks. Movements of African and Asian birds are
Woollynecked Stork 83
restricted to their respective continents and adjacent
islands, and there appears to be no interchange between the
two populations (Bannerman 1953:171).
The species is resident and locally migratory in India (S.
Ali 1964:100). Here, as in West Africa, migrations are
probably governed by the rains (Bannerman 1953:171).
C.W. Benson et al. (1971:47) reported considerable move-
ments indicative of migration in Zambia, where it is not
known to breed; in the Luangwa Valley it occurs only
during the rains from November to May. On the other
hand, western African birds move irregularly back and forth
across the equator, between the northern and southern
savannas, and avoid the rains in the wetter areas. Southern
African records suggest southward movements in the early
dry season (Apr-Jun) and northward migration in February
and March (Brown et al. 1982:182).
The Woollynecked Stork is seldom common, and gener-
ally is rare in most areas within its range. Because of its
rarity and inconspicuousness, we know little about popu-
lation levels in most regions.
In winter, we have seen as many as 30 pairs in less than
three weeks while traveling in Rajasthan and Madhya Pra-
desh, India (pers. obs. J.A.H.). It is uncommon in Gujarat,
and 8 storks and 2 nests have been reported from the Gir
Forest in Saurashtra (Luthin 1984a). During January 1989,
a total of 462 were censused in India and 21 in Sri Lanka
(D.A. Scott and P.M. Rose 1989:56). Other known popu-
lations exist in southern Vietnam (where it may breed in the
mangrove swamps), Sumatra and Sulawasi (Luthin 1987).
The total Indonesian population has been estimated at 'less
than 1000 individuals' (MJ. Silvius and W.J.M. Verheugt
pers. comm. 1989).
In Africa overall, the species is uncommon to rare, but
population numbers are probably stable (Brown et al.
1982:181). Kai Curry-Lindahl (in Brown et al. 1982:182)
has seen several hundred together in Zaire; and in Zim-
babwe, flocks of up to 200 have been reported in the Gwaai
Reserve (Smithers et al. 1957:28).
ECOLOGY
The Woollynecked Stork is a solitary species, about which
relatively little is known. It often occurs in wet grasslands or
dry ground near water, often in or near open forest. Its
overall distribution is patchy, as it occurs singly or in pairs,
often in wooded areas where marshes occur. In India, it is
'partial to water-logged ground . . . in the process of desic-
cation leaving fish and frogs high and dry' (S. Ali 1964:100).
It generally avoids deeply forested country in Africa, but it
has been seen feeding in lowland rain forest clearings in the
Congo (Curry-Lindahl in Brown et al. 1982:182). It is found
in various habitats, including grasslands, where it is par-
ticularly attracted by fires in the dry season.
In East Africa it can be found as high as 3000 m (Britton
1980:21). In India, we have generally seen this stork in
secluded forest glades and often hidden by tall marshland
reeds that surround small muddy pools. Observation
proved extremely difficult, if for no other reason than such
84 Woollynecked Stork
habitat is the preferred resting place of the tiger, which itself
discourages investigation. S. Ali and S.D. Ripley (1968:98)
considered it to be uncommon in intertidal marine habitats
in India, but this is contrary to our experience in Africa and
C.M.N. White's and M.D. Bruce's (1986:110) in Wallacea.
In Kenya, we have seen it mainly on the coast, and in
Sulawesi it occurs on sea shores (where it eats sea snakes)
and in lowland swamps and rice paddies.
In Sumatra, where it is sympatric with Storm's Stork, this
species is found mainly in open swamps, flooded grazing
land, rice fields, grasslands and dry crops (Marie and Voous
1988:66). And in Thailand, where Storm's Stork has also
bred once, the Woollynecked Stork inhabits open (logged)
semi-evergreen rain forest with grassy clearings and small
waterholes in the plains and foothills (P. Round et al.
unpub. data).
This species forages by Walking Slowly or Standing,
visually searching for prey. It feeds mainly early in the
morning or at dusk, usually singly or in pairs. It is some-
times seen in small flocks, especially outside the breeding
season. Although we once saw 16 birds foraging together in
a loose group in northern India in August. Much feeding
behaviour is reminiscent of that of the closely related White
and Black Storks. Woollynecked Storks have been seen
catching winged termites, in India, by flying back and forth
through a rising swarm and snapping up the insects (Ball, in
S. Ali and S.D. Ripley 1968:99). They sometimes attend
grass-fires, to search for injured insects and other small
animals escaping from the flames. On the Kenya coast, they
forage visually on the coral reefs, along the edge of the water
as the tide recedes, and on mudflats in tidal creeks.
The Woollynecked Stork feeds on animal matter, such as
locusts, grasshoppers, other large insects and their larvae,
water-beetles, molluscs, crabs, frogs and toads, lizards,
worms, and some fish; they probably consume some vege-
table matter, perhaps accidentally. They have been seen
eating winged termites in India (E.C.S. Baker 1929:325)
and southern Africa (P. Steyn pers. comm.), and polychaete
worms on coral reefs at the Kenya coast (pers. obs.
M.P.K.). In Sulawesi they are said to eat sea snakes, Hyd-
rophidae. They have been recorded at carrion with vultures
(Friedmann 1930:29), and often take burnt insects obtained
from grass-fires. Oil-palm fibres have been found in
stomachs, but their exact role in the birds' diet is not
understood (Brown et al. 1982:182).
This species is usually found singly, in pairs, or small
parties. Whilst McLachlan and Liversidge (1970:51) stated
that in southern Africa the Woollynecked Stork seldom
settles on trees outside the breeding season, our experience
in other parts of the range is that when disturbed it will
frequently fly to the top of a tree, choosing whenever poss-
ible a bare branch. We have watched groups of 4 or 5 birds
spend considerable time preening in tall dead tree branches.
They commonly roost in trees at night. Several authors (e.g.
A. Koenig 1910) have commented on how shy and difficult
to approach these birds are; in Zimbabwe, J.A. Scott (1975)
found them tame in the non-breeding season but 'extremely
shy and secretive when breeding'.
Predation by an immature Martial Eagle (Polemaetus
bellicosus) in Botswana was reported by P. Steyn (K.
Brouwer pers. comm. 1990).
BREEDING
This species is usually a solitary nester; small, loose colonies
of up to 4-5 nests are occasionally reported (Bouet 1955a).
The same nest may be used annually, if disturbance has not
taken place. Since birds are often seen in pairs even outside
the breeding season, it is probable that they remain paired
for more than one season, perhaps for life.
The nest is placed at the extremity of a horizontal branch
or near the top in a tall tree 10-30 m, sometimes up to 50 m,
from the ground. In India, trees such as simul (Salmalia
malabarica), neem (Azadirachta indica), or imli (Tamarindus
indicus) are used; in Africa, Brachystegia glaucescens, B. specifor-
mis, Ficus sycomorus, Acacia xanthophloea, Albizia adianthiflora or
Adansonia digitata (baobab) (S. Ali and S.D. Ripley 1968:99;
Hitchins 1974, J.A. Scott 1975, Anthony 1978b, pers. obs.
M.P.K.). Pitman (193la) and J.A. Scott (1975) reported
nests placed in trees overhanging a waterhole or pool in a
secluded section of river. Though the nest is usually not
built near other storks, if the tree is large enough other
species such as vultures (in India, Gyps bengalensis) or egrets
will often nest nearby.
In India, the nesting season varies with local rainfall
conditions (S. Ali 1953:404); most egg-laying seems to occur
during the rains. In most of the African range, on the other
hand, the species seems to be primarily a dry-season nester
(see Appendix).
Courtship displays are somewhat similar to the other
typical storks (genus Ciconia), but in well-established pairs,
which may well remain together for multiple seasons, such
displays are usually minimal (Kahl 1972c). The Up-Down
display is most similar to that of the Abdim's Stork, with the
wings held open at about 90°, displaying the brilliant orange
band of bare skin along the underside of the radius/ulna,
and the tail and undertail-coverts fanned and pumped up
and down. Rapid, wheezy bisyllabic whistles are given, and
the bill is clattered as it is raised up to about the horizontal
(Kahl 1972c).
Mackworth-Praed and Grant (1973:65) report a 'remark-
able tumbling display flight in the mating season', but
Brown et al. (1982:182) concluded that more recent obser-
vations do not confirm these reports. However, we have seen
birds in Uganda and India dive from a nest-tree and fly
about 100 m in a characteristic darting display flight, simi-
lar to one that sometimes follows the Mock Fighting display
in Abdim's Stork (Kahl 1972c and pers. obs.). J.A. Scott
(1975:205) describes a display used in response to inter-
specific disturbances at the nest that sounds similar to the
Nest-Covering display in other Ciconia storks (Kahl 1972c,
C.E. King 1988).
Both birds share in the nest-building, incubation, and the
feeding of the young (Lowther 1949:117, J.A. Scott 1975).
They build a stick nest 0.6-1.2m in diameter and about
30-60 cm deep; it has a central depression, sometimes deep
enough to hide the sitting bird, and is lined with finer twigs,
dry grass and leaves. During incubation, changeovers take

place at intervals of 2^4 h during the day. Green leaves, and
sometimes grass, are brought every few days. Repairs and
alterations to the nest are constantly carried out, especially
during the early stages of incubation.
When its mate approaches, the sitting bird will occasion-
ally become agitated and will bill clatter. When the
approaching bird lands at the nest, both birds stand, head-
bob, and clatter their bills and sometimes open their wings
in an Up-Down display (Brown et al. 1982:182). This
behaviour has not often been observed; however, most
change-overs seem to occur without display (J.A. Scott
1975), as is often the case with stork species that form long-
term pair-bonds. Such definite mating displays that have
been seen consist of allopreening by the standing bird,
directed to the top of the head and neck of the sitting
bird. Following this, both birds stand with necks bent and
heads and bills touching, circling the nest once with head-
bobbing movements repeated in unison three or four times.
The original sitting bird then sits again, and its standing
mate, with breast-feathers erect, preens both itself and its
mate, going into a half crouch and sitting on its tarsus
momentarily.
Generally 3-4, rarely 5, eggs are laid. They are rough,
white, broad to pointed ovals, and quickly become stained.
Intervals between laying are usually 2 days but sometimes
3. The incubation period is 30-31 days (J.A. Scott 1975).
Young hatch asynchronously, and the size difference be-
tween siblings may be considerable. They solicit food from
the parents with a Begging display, in which the head is
bobbed up and down and a 'filing' call is uttered (Brown et
al. 1982:183). When approached by potential predators,
such as House Crows (Corvus splendens) in India, nestlings
threaten by pointing their bills at intruders and clattering
their mandibles rapidly in a Nestling Bristle display (Kahl
1972c).
Parents feed the young by regurgitating food onto the nest
floor. A parent remains at or near the nest until the nestlings
are about 30 days old. At this time they are well feathered
and able to move about the nest. After 40 days, young move
out onto the branches and exercise their wings. They fly
after 55-65 days but remain in the vicinity of the nest for
another 3 weeks. Broods of up to 4 may be reared, although
breeding success is usually nearer to 1.5 young reared per
nest (J.A. Scott 1975, Brown et al. 1982:183).
In some areas, we have found nest observation of this
species to be difficult due to extreme shyness, and this is
confirmed by J.A. Scott (1975) who considers that behav-
iour at the nest may be adversely affected by human inter-
ference. However, in other places, e.g. several nests in India
and Uganda, birds nest near villages or houses and are not
bothered by observers nearby (pers. obs. M.P.K.).
TAXONOMY
This species was formerly placed in the monotypic genus
Dissoura, because of relatively minor anatomical characters
(Beddard 1896). It is now recombined with the other 'typi-
cal storks' in the genus Ciconia (Delacour and Mayr 1945,
Kahl 1972e). The Woollynecked Stork shows close behav-
Woollynecked Stork 85
ioural and morphological affinities with Abdim's Stork, and
its deeply forked black tail is similar to that of the Maguari
Stork of South America.
Finsch (1904), J.H. Riley (1925:28) and others dis-
tinguished birds with more red on the bill and more naked
skin on the throat as another, slightly smaller, subspecies,
neglecta. The type-locality of neglecta is in question but, as
described by Finsch (1904), its range covers Celebes
(=Sulawesi), Java, Sumbawa, Lombok and the Philippines.
Chasen (1935) gave its type location as Java, but C.M.N.
White (1974) considered that he erred. E.C.S. Baker
(1929:325) was unable to separate neglecta from the main-
land populations; he stated that the amount of naked skin
on the throat varies greatly according to age, with older
birds having more extensive areas of bare skin. Whilst most
birds on the Asian mainland are reported as having a black
bill with red on the culmen and at the tips of both man-
dibles, this can not be taken as unalterable, and the bill
may, like that of other species, vary according to season and
maturity. Examination of museum skins has shown that,
while specimens from the Sunda Islands do show more red
in the bill than those from India, many African birds have
nearly as much red (pers. obs. M.P.K.). The differences are
in our opinion too few and inconsistent to recognize neglecta
as a valid race. A further discussion of this problem is found
in C.M.N. White and M.D. Bruce (1986:110-111).
Previously, many ornithologists considered birds we now
recognize as Ciconia stormi to be a subspecies of C. episcopus.
The recent discovery of the Storm's Stork breeding in
southern Thailand (Nakhasathien 1987) and southeastern
Sumatra (F. Danielsen et al. pers. comm. 1990)—where it is
sympatric with C. episcopus without interbreeding—has
clearly shown to us that it is a separate species (see Storm's
Stork).
CONSERVATION
Once widespread, this species has suffered overall popu-
lation reductions from habitat destruction and shooting
(Henry 1971:381). Although it remains uncommon but
stable in Africa, the widely dispersed populations of Asia
require urgent conservation measures in the establishment
of reserves. The Melaleuca forests in Vietnam, where the
storks breed, are fully protected (Luthin 1987). The limited
information on this species makes more a detail conser-
vation programme difficult. Thus range-wide surveys are
needed to determine population size and status in both
Africa and Asia. These data could lead to the development
of appropriately situated reserves.
Like other solitary nesting storks, which are likely to pair
for life, this species returns regularly to traditional nesting
sites, which are usually in a large tree. Without protection of
these trees, a number of which are well known, this species
will undoubtedly suffer further decline.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
pages 292-293.
 Storm's Stork
Ciconia stormi (Blasius)

Melanopelargus episcopus stormi Blasius, 1896, Mitteil. Geogr. Gesell. Naturhist. Mus. Liibeck, ser. II
pts 10-11, p. 120: western Borneo, Pontianak

Other names:
Borneo wolhalsooievaar (Dutch); Nok kra su-um (Thai); Burung ginti (Pasir Panjang/Kalimantan)

IDENTIFICATION
A largely black medium-sized stork, Storm's Stork stands 85 cm tall, with white on the hindneck,
abdomen, and under tail. It is similar to, but slightly smaller than, the closely related Woollynecked
Stork.
The adult's colour and pattern are very similar to the Woollynecked Stork including a black cap,
downy white sides and back of neck, dark body plumage, and white belly and undertail coverts. Unlike
the Woollynecked, which has an entirely white neck, black feathering in the Storm's Stork projects up
the front and sides of its white neck in two wedge-shaped patches. As a result, the only white on the
head and neck is restricted to the lower cheeks and nape and to a very narrow wedge extending down
the midline of the foreneck for the upper third of its length (Ogilvie-Grant 1903, Nakhasathien 1987).
The black cap has a definitive edge where it meets the neck, as in the Asian race of the Woollynecked,
but it is a bit more extensive, coming forward to the bill and further back on the nape (pers. obs.
M.P.K.). The black tail is deeply forked contrasting with the longer white undertail-coverts. [Some
confusion exists in the literature about which actual tail-feathers and undertail-coverts are black and
which are white. D.R. Wells (pers. comm. 1990) and C. King (pers. comm. 1990) confirmed in museum
specimens that, as in the Woollynecked, all the feathers of the deeply forked tail are black and all the
undertail-coverts are white.]
The bill is vermillion, often with some pinkish red-orange on the sides and a yellow tip. A substantial
area of facial skin is a dull orange to orange-yellow or golden. Particularly below the eye, this naked,
golden skin appears slightly swollen (C. King pers. comm.). On some but not all individuals, the
culmen is slightly concave (i.e. recurved) and there is a small knob where the culmen meets the forehead
(Ogilvie-Grant 1903, Medway and Wells 1964, 1976:95). The legs and feet in the adult are dull red,
with darker grey joints, but often become covered in extreta turning them a paler colour. The iris is red.
There is an excellent coloured plate of this species, accompanying the article by Ogilvie-Grant
(1903:pl.V).
A captive Storm's Stork at Walsrode, Germany, acquired as a juvenile in early 1988, had in May
1989 a stripe of bright orange, naked skin down the under side of the ulna, visible when the wing was
opened (pers. obs. M.P.K.). In this it is similar to C. episcopus.
The sexes are similar, although, as in other storks, the male is probably slightly larger. This needs to
be confirmed. At a nest in Sumatra, one of the adults was slightly larger and more glossy on the breast,
throat, and back; this was assumed to be the male (F. Danielsen et al. pers. comm. 1990).
The bill and soft parts brighten during the breeding season. The bill becomes vermillion to orange-
red. The throat, cheeks and part of the facial skin orange grading to pinkish-orange near the neck
feathers. A large area around the eyes is yellowish-golden. An adult in full breeding plumage is well
illustrated accompanying the articles by Nakhasathien (1987) and Round et al. (1988).
At 1-3 days old, young are covered in white down, with a black bill and black crown. In less than a
month, they roughly double in size, and black feathers start emerging on their throats, wings and
88 Storm's Stork
bodies. Their bill is a blackish-grey with a yellow-orange
tip. The facial skin starts a pale yellow and changes to dark
grey; the gular pouch is a brighter yellow, with small black
spots. The iris of nestlings is brown. Their legs start as a
light yellow, gradually changing to pink with darker joints.
The areas covered by black feathering increase greatly after
30 days. By about 45 days, the nestlings resemble the adults
in feather pattern, but they were smaller and had shorter
bills; they were fully feathered by 60 days. Unlike the adult,
the black feathering on the head extends below the eye to
the base of the bill. The legs are dark greyish flesh initially,
but the tarsus turned a dull reddish-orange at about
3 weeks. Captive nestlings in Thailand could fly by 90 days,
but wild young in Sumatra left the nest at about 60 days
(Nakhasathien 1987, Danielsen et al. pers. comm. 1990).
Captive juveniles have browner and duller body plumage
than adults. They show a slightly more densely feathered
upper neck, and the pinkish orange area of facial skin is a
less intense dull gold. Their bills are a lighter red than
adults (C. King pers. comm. 1990).
A bird we observed in captivity at Walsrode, Germany,
on 10 May 1989, gave a short series of sibilant whistles, very
similar to those heard from the Woollynecked and Abdim's
storks; they were not loud and were just audible at 15 m
distance (pers. comm. M.P.K.). Elsewhere the adult's voice
is described as 'Karau' (Smythies 1981:30) and that of the
nestlings as a 'loud harsh Krack, krack, krack' (Nakhasath-
ien 1987). Begging calls of the young have also been charac-
terized as a froglike 'Skhrekeh . . . keh . . . keh . .. keh,
skhrekeh . . . keh . . . keh* (F. Danielsen et al. pers. comm.
1990).
This species is often seen soaring high over rivers or forest
clearings (Nash and Nash 1988). As with other stork
species, soaring behaviour is apparently very 'contagious',
for when one bird began to soar near the nest-site in Suma-
tra, several others came from different directions to join in,
until there were 6 birds soaring together for 30-60 min (F.
Danielsen et al. pers. comm. 1990).
Storm's Stork is slightly smaller than the Woollynecked
Stork, with which it might occur in Sumatra or southern
Thailand. When viewed from below in flight, the neck of the
Storm's appears wholly dark (Holmes 1977). Although
there is probably some overlap where habitats merge, the
two species are ecologically segregated, with Storm's Stork
generally preferring denser, undisturbed forested areas and
the Woollynecked occurring in more open country and more
often in disturbed habitat (Nakhasathien 1987).
DISTRIBUTION AND POPULATION
This species is resident in Borneo and southeastern Suma-
tra, and a rare resident or irregular visitor in peninsular
Malaysia and Thailand (Collar and Andrew 1988:15). It
has recently (Nakhasathien 1987) been discovered nesting
in southern Thailand and in South Sumatra (F. Danielsen
et al. pers. comm. 1990). The majority of recent sightings
have come from Borneo, where it is uncommon but wide-
spread (Smythies 1981:30; Luthin 1987). In recent years,
observations of individuals or small groups have been
reported from Central Kalimantan, East Kalimantan, east-
ern Sarawak, Brunei and eastern Sabah (MacKinnon 1983,
Luthin 1987). In Sumatra and neighbouring islands, there
are a number of sightings. A specimen was collected from
eastern Sumatra, and another from North Pagai, West
Sumatran Islands. There are several records in peninsular
Malaysia, including Sungei Tekam Forest Reserve and
Kuala Lompat (Medway and Wells 1976:95) and, more
recently, in Taman Negara and on the Pahang-Johore bor-
der (G.W.H. Davison and D.R. Wells pers. comm. 1990).
The discovery, in 1986, of a nest of Storm's Stork in
southern Thailand extends the range of this species over
500 km further to the north (Nakhasathien 1987).
Nothing is known about migration. Probably no real
migration as such occurs but storks most likely wander in
response to local conditions (e.g. drought, disturbance,
changes in food supply). Transient birds have been reported
almost 20 km from the nearest typical habitat in Sumatra
(F. Danielsen et al. pers. comm. 1990).
This is a very rare to uncommon species, many obser-
vations being of only single individuals or pairs. Never an
abundant species, there has apparently been a decline in
numbers and it is now considered rare everywhere (W.B.
King 1978). It is distinctly local in the heart of its range in
Borneo, where it is thought to be decreasing. In western
Malaysia, recent records include a bird seen in the Sungei
Tekam Forest Reserve, Pahang (February 1969) and 2 near
Kuala Lompat, Pahang, in the Kerau Game Reserve
(February 1970) (Medway and Wells 1976:95). Since then
one or two birds have been reported each year (Wells pers.
comm. 1990). An unusual concentration of 12 individuals
was seen in Brunei (Holmes 1969 cited in Smythies

1981:30), and 6 (possibly as many as 12) were seen, in
groups of 1-3, in East Kalimantan along the Telen River
(Holmes and Burton 1987; D. Holmes pers. comm. 1990).
Up to 7 individuals have been seen near the Sembilang
River, in South Sumatra (F. Danielsen pers. comm. 1990).
The recent northward extension of its known range into
southern Thailand is based on a single known nest. As it
was found in an area of forest which is now flooded by dam
construction, the small population may well have perished
even at the point of its discovery (Nakhasathien 1987, P.
Round et al. unpub. report). A recent estimate of the total
population in Indonesia (i.e. Kalimantan and Sumatra)
stated that they 'may well number less than 300' (MJ.
Silvius and W.J.M. Verheugt pers. comm. 1989), although
M. Silvius (pers. comm. 1990) now considers that estimate
'rather too high'. In total numbers this is probably one of
the least numerous of all the storks in the world.
ECOLOGY
Storm's Stork is a solitary species of dense primary forest
and the swamps, rivers, and small streams running through
it. Most recent sightings in Borneo were along rivers adjac-
ent to dense forest. In Borneo, Smythies (1981:30) reported
it frequenting open, grassy, freshwater swamps subject to
periodical flooding, river banks in remote forest, and even
paddy-fields. In Sumatra it frequents dense primary and
logged riverine and swampy lowland forest (Marie and
Voous 1988:66) and has been seen feeding in small forest
ponds (M. Silvius pers. comm. 1990). In the Malay Penin-
sula it frequents rivers and damp clearings in lowland forest
(Medway and Wells 1976:95). The nesting habitat in
southern Thailand was thick lowland forest with very large
trees, such as Hopea ferrea, Cynometra bijuga, Dipterocarpus
spp., Mesua ferrea and Ficus spp. Trees exceeding 10 cm
(diameter at breast height) had a density of roughly 500-
800 trees per hectare and there was an 80-90% canopy
cover (Nakhasathien 1987). The undergrowth was mainly
rattans and other palms, bamboos, shrubs and climbers,
and this correlates with similar lowland forest regions in
Borneo. In general this species seems to be less tolerant of
disturbed habitat than is the Woollynecked Stork (MJ.
Silvius and W.J.M. Verheugt pers. comm. 1989).
Little is known of feeding behaviour. Birds forage along
the mudbanks of main rivers in central Kalimantan (Nash
and Nash 1988). Its local name in Thailand ('nok kra su-
um') refers to the birds' way of fishing, by quiet stalking,
along the bank of a stream in dense forest (Nakhasathien
1987). Adults from the Sumatra nest foraged mainly in
primary swamp forest 2-3 km from the nest (F. Danielsen et
al. pers. comm. 1990). They were most active away from the
nest in the early mornings. D. Holmes (pers. comm. 1990)
has seen them feeding in Lampung, Sumatra, in a small
wetland within a dryland secondary forest.
Similarly, little is known of the diet of Storm's Stork. Food
items brought to the young by parents at the Sumatra nest
were mostly fish, 5-7 cm long and weighing an estimated
10-30 g each, also some frogs, and worms 10-15 cm long (F.
Danielsen et al. pers. comm. 1990). It is reported that it eats
Storm's Stork 89
earthworms in Borneo (Smythies 1981:30). Other food is
probably similar to that of the Woollynecked Stork, but
more data are needed.
BREEDING
The discovery of a nest in southern Thailand, by Seub
Nakhasathien on 27 September 1986, was the first nest of
this species known to science. The nest was 19 m up in the
fork of a 27m tall dipterocarp tree (Dipterocarpus baudii) on
the bank of the Khlong Mon, a tributary of the Khlong Phra
Saeng, at 69m above sea level. This is in Khlong Saeng
Wildlife Sanctuary, Surat Thani Province, Thailand, at
9°05'N 98°30'E. The nest was a flat platform 15cm deep,
with an external diameter of 50 cm. It was constructed of
dry sticks 15-60 cm in length and lined with dry leaves and
some down. A second, unoccupied, platform was found
some 200 m away in the top of another 30 m tall diptero-
carp. A local man reported that before leaving the area in
1985, he had seen 4 birds feeding along a nearby tributary of
the Khlong Mon (Nakhasathien 1987).
On 26 April 1989 a half-built nest, the second known, was
found in the Sembilang area of South Sumatra. This site
was on the bend of a river 8-12m wide, about 20km
upstream from the sea, near the boundary between man-
grove and freshwater swamp forest; within 5 km of the nest,
82% of the area was covered with primary swamp forest and
17% by mangrove forest, as determined from satellite pho-
tos. The nest was 8.3 m above the river on the second lateral
branch of a 18m high Rhizophora mucronata tree. The nest-
platform, when complete, was 30-40 cm in diameter and
10-15 cm in height. It was constructed mainly of branches
of Rhizophora mucronata 15-25 cm long and 0.5-1.0 cm thick,
and lined with leaves ofXylocarpus granatum and draped with
epiphytes (F. Danielsen et al. pers. comm. 1990). [It is not
known whether the birds nested again, at the same site, in
1990; the area could not be checked because of budget
restraints (F. Danielsen pers. comm. 1990).]
Practically nothing is known of courtship behaviour. It is
probably similar to that of the closely related Woollynecked
Stork. At the Sumatra nest, very little in the way of ritual-
ized displays was seen; when one bird returned to the nest to
feed the young, the other adult usually flew away (F.
Danielsen et al. pers. comm. 1990). If crows, monkeys, or
raptors came near the nest, the adult spreads its wings to
cover the nestlings, in what sounds much like the Nest-
Covering display reported in the Maguari Stork (Kahl
1972c).
A bird we observed in captivity at Walsrode, Germany,
on 10 May 1989 (pers. obs. M.P.K.), gave a low-intensity
form of the Up-Down display. Initially it raised its bill to
about horizontal, then it clattered rapidly but softly 15-20
times, while raising its bill further to about 20° above the
horizontal. The entire performance was reminiscent of that
seen in the Woollynecked Stork (Kahl 1972c).
Very little is known about behaviour at the nest. When the
Thai nest was discovered on 27 September, the adults were
apparently incubating. Two chicks hatched about 21-24
October, and when 1-3 days old were seen to crouch down
90 Storm's Stork
and remain motionless when the loud calls of a Crested
Serpent Eagle (Spilornis cheela) were heard. By 22 November
the nest had been destroyed and the young stolen by a local
villager, who dislodged the nest and chicks with a long
bamboo pole. The stolen young from the nest were located
and delivered, for captive rearing, to the Nature and Wildlife
Educational Centre at nearby Surat Thani. Here they were
reared on freshwater fish, cut into small pieces. The newly
constructed dam having now completely flooded this area of
lowland forest, any remaining Storm's Storks must be con-
sidered lost to the area, and possibly to Thailand.
In the 1989 Sumatra nest, an unhatched egg was found.
When taken from the nest, it was 'olivebrown with black
and greyish black areas'; however, after cleaning, it was
'basically white in colour' (F. Danielsen et al. pers. comm.
1990). This egg, apparently the only one of this species
known, is preserved at the Zoological Museum, Bogor,
Indonesia.
When humans approached, the adult at the Sumatra nest
crouched low on the platform, until only a small mass of
black feathers, 'with only the eyes free to follow the
intruder', was visible from below (F. Danielsen et al. pers.
comm. 1990).
Adults brought food to the young about 5 times/day dur-
ing their first month and about 2 times/day during their
second month. Food was regurgitated onto the nest-floor
and picked up by the nestlings. Mostly small fish, plus some
frogs and worms (still alive when presented) were fed to the
young (F. Danielsen et al. pers. comm. 1990).
While the young were in the nest, both adults returned to
sleep in or near the nest each evening.
TAXONOMY
Previously the Storm's Stork has frequently been considered
a race of Ciconia episcopus (Kahl 1972e, 1979b). It is clear
now that both species occur, or have occurred, within the
same areas. Holmes (1977) and Silvius and Verheugt (1986)
gave evidence for the occurrence of both Storm's and
Woollynecked Storks in southeastern Sumatra, and Storm's
has now been found nesting there (F. Danielson et al. pers.
comm. 1990). In Thailand, the discovery of nesting Storm's
Storks (Nakhasathien 1987) establishes its presence within
the former range of the Woollynecked Stork there.
Unlike the putative race C. episcopus neglecta, C. stormi is
quite distinct from C. episcopus. It has a striking orange and
yellow facial skin coloration, which is apparent from the
nestling stage onward, and a different feather-pattern on the
sides of its neck. The habitat preferences of the two popu-
lations are different, with Storm's keeping to more heavily
forested areas. On the basis of available information, espe-
cially the plumage and soft part differences, ecological
differences, and its wide zone of sympatry with C.e. episcopus,
we have concluded that Ciconia stormi should be considered a
separate species.
CONSERVATION
This lowland forest bird is clearly at risk throughout its
range, due to forest destruction and hunting. Although it is
listed as a protected species in Indonesia, Sarawak, and
Thailand, such designation has brought no real protection
or conservation action.
The saga of the discovery of the first Storm's Stork nest
ever to be found (Nakhasathien 1987), its destruction by
local hunters, its flooding by dam construction, and the
likelihood of this species' complete extirpation from Thai-
land at the very moment of its discovery, presents vividly
the situation as it exists in Asia today. There appears to be
little or no appreciation of the needs of conservation in many
areas. Nakhasathien (1987) stated that the impact of the
Chiew Larn Dam upon lowland bird communities was
never addressed. In what was supposed to be an environ-
mental impact assessment before the dam was approved, it
was stated (Anon. 1980 cited in Nakhasathien 1987) that 'in
regards to birds . . . the adverse effects of the impoundment
are likely to be minor, because most birds are mobile and able to
flee away in the event offloading [our emphasis]. It would seem
difficult to find a statement that more clearly indicates the
need to explain the effects of such developments to govern-
ment bodies unfamiliar with conservation issues.
W.B. King (1981) stated that 'the extensive destruction of
forests in Borneo is believed a major factor in the decline.'
Such widespread lowland forest destruction occurs through-
out Southeast Asia. The only hope for the survival of species
like Storm's Stork is for areas of complete protection to be
established, not only against the destruction of the trees for
commercial use or for dam construction but also against local
hunters. Smythies (1981:30) stated that in Borneo, Storm's
Stork is 'probably decreasing because it is shot for food.' In
many cases the hunters are not original forest dwellers by
tradition but have been attracted to such areas either for
exploitation of the land or water, under their own volition or
by government edict. Although direct persecution kills some
birds, habitat destruction is by far the greater danger.
Range-wide surveys are needed to locate the species and
assess its breeding and feeding requirements, followed by its
protection on large tracts of land. [According to F. Daniel-
sen (pers. comm. 1990), the best time to locate this shy
species is during early mornings when it is most active.]
Clearly, with so little known about its biology and ecology,
much basic study is required. The most critical long-term
threats appear to be the Indonesian Government-
encouraged movement of people into the storks' forest habi-
tat, and unsuitable forestry practices in swamp forest where
regeneration is poor.
Capture, mainly of nestlings, for supply to the inter-
national zoo trade has apparently increased in recent years.
Storm's Storks have recently appeared in zoos in Kuala
Lumpur, Malaysia and Walsrode, Germany. The price of
the species in the bird markets of Singapore has doubled
(from S$300 to S$600) in the past year (F. Danielsen et al.
pers. comm. 1990). Such trade, if not restrained, could put
fatal pressure on such a rare and localized species.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 294.
 Maguari Stork
Ciconia maguari (Gmelin)

Ardea Maguari Gmelin, 1789, Syst. Nat., 1(2), p. 623; based on 'Maguari' of Marcgrave, 1648, Hist.
Rerum Nat. Brasiliae, p. 204: northeastern Brazil

Other names:
American Stork (English); Cigiiena comun, Cigiiena Americana, Maguari tabuyuyi, Magoray,
Yiulo, Gulo, Cabeza de Heuso, Sa pyta, Baguari Pillo, Gaban Peonio, Tuyango, Malvinero
(Spanish); Cauaua, Cegonha, Joao-grande (Portuguese); Erie, Reddie foetoe (Dutch); Pillo, Pillu
(Araucano); Tuyuyu (Guarani)

IDENTIFICATION
The Maguari is a large stork, standing 97-120 cm tall. It has a straight bill and is largely white, with
black flight feathers in the wing and a forked black tail.
The glossy black of the adult's wing involves most of the feathers, including scapulars, greater coverts
and flight feathers. The forked black tail—created by the outer tail feathers being longer than the
inner—is shorter than the stiff white undertail-coverts (well illustrated in Chubb 1916:156), which may
function aerodynamically. Nonetheless, the black tail can be seen through the under-coverts in
overhead flight (Wetmore 1926:62).
The bill is mainly bluish grey bordered with red, shading to dark maroon and blackish in its distal
third. It is long, relatively thin and, except for its thicker base, markedly heron-like. In fact, in its
original scientific descriptions, the bird was confused with a heron (Molina 1782). The head and neck
feathers are long semi-plumes, which are erectile and function in display. The throat and pebbly-
textured lores and orbital-skin are orange-red, turning redder during courtship. The legs are purplish
red, and the iris pale lemon-yellow to cream-white (Wetmore 1926:63, Sick 1985, B.T. Thomas 1988).
The sexes are similar, although males are slightly larger and have a more recurved bill (Kahl 1971e).
During courtship the plumage is bright from a fresh moult, the base of the bill is greenish grey
whereas the tip is violet, with a diffuse zone of whitish violet between them (S. Narosky pers. comm.
1990).
At hatching, nestlings weigh 76-90 g, and their sparse down is 'snow-white' (B.T. Thomas pers.
comm. 1990). The bill is dull grey, the gape pink, the gular area whitish yellow, the legs and feet pink,
and the rest of the skin bluish grey. By 8-10 days the body down is greyer, the head-down being curled
and blackish; the nestling's bill, legs and feet become shiny black. The unfeathered throat is orange, and
there is also a pale yellow stripe extending up the ventrum. This iris is dark brown. Succeeding the
whitish down present at hatching, the young have two coats of very dark feathers, which together last
for much of the nestling period (c. 10-75 days). By 3 weeks it may fade to olive-brown (Kahl 1971e, B.T.
Thomas 1979b, 1984).
By about 8 weeks, the juvenile plumage begins to appear. Development of this white plumage
continues during the remainder of the first 3 months. By fledging, the gular patch has turned to pale
scarlet. By 3 months, the legs and feet start to turn pinkish, the bill becomes bi-coloured, and the
plumage resembles that of adults. The skin around the eye remains black for about a year, before
turning red, and the dark brown iris does not begin to turn to yellowish until the second year. In fact,
the dark iris is the best field mark for distinguishing juvenile Maguari Storks from adults (B.T. Thomas
1984).
Nestlings of no other stork species have dark coloured down. Black nestling plumage may function to
make young in terrestrial nests less visible to predators (Fernandez 1920, Kahl 1971e, B.T. Thomas

1984, C.E. King 1988), or it might aid thermoregulation in
cold weather by absorbing solar radiation (Cowles et al.
1967). Adults give wheezy, hissing, bisyllabic whistles
during the Up-Down display; these are hoarser, lower-
pitched and slower than those given by other species of
Ciconia (Kahl 1971e, 1972c). During some social displays,
especially hostile ones, they loudly clatter their bills (B.T.
Thomas 1986, C.E. King 1988).
Young beg with a 'Ehehe-ehehe' call. A variety of other
sounds made by nestlings is described by B.T. Thomas
(1984).
The Maguari flies with neck and legs outstretched with
strong wing beats alternating with gliding. It soars regularly
in Buenos Aires Province, Argentina (Gibson 1880, Weller
1967) and in the Venezuelan Llanos (B.T. Thomas pers.
comm. 1990). The flapping rate in normal flight averages
181/min( B = 32) (Kahl 1971b).
The Maguari Stork is distinguishable from herons by its
white colour, large grey and maroon bill, and black wings,
and in flight by its outstretched neck. The two other large
sympatric storks have featherless black heads; the Wood
Stork is smaller and the Jabiru is larger than the Maguari.
DISTRIBUTION AND POPULATION
The Maguari is found in South America, primarily in low-
land areas east of the Andes, from eastern Colombia,
through inland Venezuela, and the Guianas (sparsely), Bra-
zil (mainly south of the Amazon), eastern Bolivia, south-
ward through Paraguay, Uruguay, and Argentina to Chu-
but Province. It is rare, and probably non-breeding, west of
Maguari Stork 93
the Andes in Chile, where it has occurred as far south as
Magallanes Province (A.W.Johnson 1965:150).
The species is at least partially migratory, although little
is known of its movements. Birds leave the Venezuelan
Llanos when the marshes dry and the aquatic food disap-
pears (B.T. Thomas 1985). It also occurs only seasonally in
Bolivia and northern Argentina. Occurrences further south
and in Chile may be related to drought dispersal (A.W.
Johnson 1965:150). In Surinam, it is a rare visitor to coastal
areas from March to May (Spaans 1976a). B.T. Thomas
(1986) observed that several months after leaving the nest,
the young left the nesting area, and none of the 128 ringed
birds were seen again in the area until they returned to
breed at 3-4 years of age.
Apparently widespread in typical habitat over much of its
range, this species is found nowhere:in large numbers (B.T.
Thomas 1987). Although quantification is lacking, it
appears to be most numerous in the Pantanal and Chaco of
Argentina, Brazil and Paraguay (Short 1975, Nores and
Yzurieta 1980). Brooks (1991) found it to be relatively
common in the Paraguay Chaco, at a density of one stork
per 1.16 km2. Populations appear larger in Amazonas and
Rio Grande do Sul, Brazil (Sick 1985). Surveys in the
Llanos of Venezuela found 82 individuals in 1983 (Luthin
1984a); however, studies there have shown that it has de-
creased markedly within the past decade (B.T. Thomas
1986, 1987).
ECOLOGY
Preferring shallow marshes, mudflats, tropical wet savan-
nas, the Maguari Stork is a bird of open country and
generally stays away from forested regions (Wetmore
1926:62). Its usual foraging site is very shallowly flooded
wet meadow, in water less than 12 cm deep and more rarely
in water as much as 30cm deep. It feeds solitarily or in
pairs, especially during the nesting season, but will feed in
larger aggregations and with other species of wading birds,
especially outside the nesting season, Hudson (1920:114)
reported that in the evening they gather in large numbers
around water but in the daytime disperse over the feeding
grounds. Primarily a lowland species, it occurs occasionally
in the foothills of the Andes (e.g. at Cochabamba, Bolivia—
Remsenetal. 1985).
The Maguari Stork feeds visually by Walking Slowly,
usually in shallow (2-30 cm) water, with its neck extended
and partially open bill near the water, Pecking and Grab-
bing prey items. It may search carefully in dry fields and has
been observed to lift up pieces of cow-dung to inspect
underneath (B.T. Thomas 1986). It also will Wing Flash
and Run short distances after prey. B.T. Thomas (1986)
described observations of 'aerial pawing', a behaviour that
would appear to be Foot Stirring in the air rather than in the
water. Maguari Storks will sometimes Grope in the water or
Probe in mud. When they catch a large prey item, they bite
it a few times before swallowing. They have been observed
foraging at grass-fires in Paraguay (Kahl 1972c).
The diet is quite broad, including earthworms, insects,
insect larvae, freshwater crabs, frogs, tadpoles, snakes, fish
94 Maguari Stork
and eels, small mammals, and eggs and young of marsh
birds (L. Fraser 1843, Pelzeln 1868-71:304, Gibson 1919,
Fernandez 1920, Zotta 1940, Kahl 197le, B.T. Thomas
1984, 1985). A large rail was found in the gullet of a bird
shot in Patagonia (Bent 1926:70), and a Maguari was once
observed eating cow-dung in Venezuela (B.T. Thomas
1985).
In the Llanos of Venezuela, of 344 food items eaten by
adults or brought to the young, 44% of the diet was fish and
eels, 32% frogs and tadpoles, 12% rats, and 12% other
invertebrates and vertebrates (B.T. Thomas 1985). Food
brought to young differs in different years, presumably
depending on availability.
BREEDING
In Venezuela, storks return to their breeding areas in flocks
and gather in groups on feeding and loafing grounds. Most
of the time in these parties is spent standing or preening.
Some pairs reform in successive years on the same nest-site
or in the same area (B.T. Thomas 1986). Sites may be
reused, but most nests are rebuilt each year, as they do not
usually last between seasons.
Nesting generally starts early in the rainy season, so that
young fledge at the end of the rains (B.T. Thomas 1987).
However, it differs in timing throughout the storks range,
and to some extent from year to year. Egg-laying occurred
between May and November in different years in the Llanos
of Venezuela (B.T. Thomas 1984), depending on variations
in the beginning of the rainy season. Within an area in a
given year, egg laying is fairly synchronized (B.T. Thomas
1985, 1986). In Argentina, nesting occurs in August and
September (S. Narosky pers. comm. 1990).
In the northern part of its range (e.g. Venezuela), the
Maguari nests in small trees and bushes, usually less than
6 m high; nests in dead palms tend to fail because the trees
break off or young fall out (B.T. Thomas 1984). In other
areas (e.g. eastern Brazil, Argentina), it nests on the ground
in marshes (Hagmann 1907, Kahl 1971e). Birds may nest
singly, semi-colonially in well-dispersed groups, near colon-
ies of other species, or in colonies of 5-15 nests; in colonial
situations, nests may be within 50 cm of each other but are
separated vertically in the tree or bush (B.T. Thomas 1984,
1986). In Venezuela, 77% of nesting was colonial, and
colonial nesters were more successful than solitary nesters
(B.T.Thomas 1984, 1985).
Pair-formation and the reunion of pairs from previous
years, which appears likely, may take place in assemblies on
the ground away from the nest-site, as birds display to each
other by walking about stiff-legged and giving Up-Down
displays (B.T. Thomas 1986). This appears to be in contrast
to pair-formation in other species of storks, which takes
place in trees at or near the nest-site; however, in the
Maguari such aberrant pair-formation may reflect an adap-
tation to ground-nesting, which these birds do over much of
their range.
Thomas' observations were made in the Llanos of Vene-
zuela, where the storks are tree-nesters and primarily col-
onial; nothing is known of pair-formation in the remainder
of the range, where they nest mainly on the ground and
mostly solitarily or semi-colonially. The elongated feathers
on the lower foreneck are strongly erected in many displays.
A variety of ritualized breeding displays were described
by Kahl (1972c) and C.E. King (1988). Many of the dis-
plays are similar to those seen in other species of 'typical
storks'. The Head-Shaking Crouch, a display which appears
characteristic of and confined to storks of the genus Ciconia,
has not been observed in the Maguari, although it may be
expected to occur.
The Up-Down displays observed in Argentina involved
the displaying bird throwing its head to near the vertical,
clattering the bill loudly and slowly 6-10 times; this was
preceded and/or followed by 4-6 wheezy, two-part whistles
(Kahl 1972c). B.T. Thomas (pers. comm. 1990) described
the vocalization as a throaty hiss.
During Up-Downs observed in captives, C.E. King (pers.
comm. 1990) recorded that birds stood erect with the body
nearly vertical, the tail lowered and the wings held close to
the body. The head was thrown back, but not so far as in the
Up-Downs of White and Oriental White Storks. Males
tended to throw the head further back than females. The
bird then bent forward, brushing the bill through the
erected neck feathers, and the display terminated with the
bill near or touching the ground. The wheezy vocalizations
were given during the 'back' portion of the display and
before the commencement of the bow.
Up-Downs observed in Venezuela were rarely associated
with bill-clattering (B.T. Thomas pers. comm. 1990). And
C.E. King (1988, pers. comm. 1990) did not observe clatter-
ing in over 562 Up-Downs given by 10 captive birds from
Paraguay and Argentina. It would appear that clattering
during the Up-Down is rare, perhaps absent in some birds,
in the Maguari Stork. This is similar to the situation with
the Black Stork, in Eurasia and southern Africa. After nests
were constructed, the storks continued to give Up-Down
displays, which lasted an average of 9.25 s (n= 7) (B.T.
Thomas 1986).
Bill-clattering in Venezuela was observed to be associated
with instances of aggression, threat and annoyance (B.T.
Thomas pers. comm. 1990). It was given when humans or
other birds, such as the Great White Egret (Egretta alba),
Barefaced Ibis or Carib Grackle (Quiscalus lugubis).
Copulation occurs on the nest, often after presentation of
grass by the male. The Copulation Clattering display in-
cludes the members of the pair snapping their bills together
or the male biting in the neck feathers of the female; 13
successful copulations were measured as lasting an average
of 8.8s (B.T. Thomas 1986).
Both B.T. Thomas (1986) and C.E. King (1988) observed
courting birds carrying grass or other material in their bills,
and this may have display function. In nest-building, males
bring most of the sticks and females do most of the construc-
tion, although both do participate in both jobs. Nesting
material, especially the lining, is added throughout the
breeding season (B.T. Thomas 1986).
The nests of the Maguari are unusual among storks in the
extensive use of grass or reeds. Marsh nests may be made
entirely of reeds (Cyperus giganteus), grasses and other aqua-

tic plants [Polygonaceae, Solinaceae (S. Narosky pers.
comm. 1990)]. They look very much like crane nests. These
nests are large conical structures, which can be up to 2.5 m
wide and 1 m tall, built up from the floor of the marsh rising
to 0.5 m above the water (Hagmann 1907, Fernandez 1920,
Kahl 1971e). S. Narosky (pers. comm. 1990) found nests
80 X 150 cm in external diameter. We found similar ground
nests in tree islands. Arboreal nests, too, may be made
entirely of grass but most are made of sticks (<1 m
long X <2cm diameter), covered with wet grass that is
poked into the nest structure and hardens as it dries.
The eggs are nearly smooth, dull white, and oval to
subelliptical; the yolk is flame scarlet in colour. The clutch
size is 2-4 eggs, averaging 3.2 in 49 Venezuelan nests (1974—
76); smaller clutches may occur in dry years. Eggs are laid
on alternate days and incubation begins with the second or
third egg. Egg-laying is highly synchronized in closely
adjacent nests. Both adults participate in incubation, which
takes 29-32 days. Young may hatch as much as a week
apart and it is common to find weight differences of 500-
1400 g between siblings (B.T. Thomas 1984, 1986).
Young 6-7 days old emit a guttural call. They give Up-
Down greeting displays—with rapid, but largely inaudible,
bill-clattering—to returning adults and then drop to their
tarsi and commence the Begging display (Kahl 1972c). Both
adults feed the young, with undigested food regurgitated
onto the nest floor. The size and quantity of food brought to
nestlings increases with their age. An 18-day old nestling
was given an eel 60 cm long and weighing 326 g; it took
several hours to swallow (B.T. Thomas 1984). Parents are
occasionally observed giving water to nestlings at midday,
and they often shaded eggs or nestlings, with their backs to
the sun and wings partially spread (B.T. Thomas 1986).
One parent usually remains with the young at all times until
they are about 6 weeks old. After the age of 25-35 days, but
well before they can fly, young in terrestrial nests often
wander off the nest, into the adjacent marsh (Kahl 1972c);
however, young in elevated nests are 'nest-bound' until they
can fly (B.T. Thomas 1984). The nest and its contents are
defended by both parents, who give Arching displays, Aerial
Clattering and Nest-Covering displays in response to poten-
tial predators near the nest. At solitary nests, adults perform
a Lead-Away display, in which they approach a human
intruder and clatter their bill, then walk away slowly, lead-
ing them from the nest (B.T. Thomas 1984, 1985). Feath-
ered young also defend their nest, emitting a grunting or
bleating cry and rattling their bills (Centro Editor 1983b, S.
Narosky pers. comm. 1990).
Young remain in the nest for 60-72 days and weigh about
4 kg when they leave the nest. Upon fledging, young form
groups with other young at feeding areas. Juveniles appar-
ently can recognize their parents individually, and they
emerge from these groups to be fed by them, either in the
marsh or back at the natal nest, for 5-6 weeks after fledging.
Juveniles also return to their nest to roost at night for up to
6 weeks after fledging. There is little sibling rivalry, and
success from 55 nests over 9 years in Venezuela was 52%
(i.e. 89 young fledged from 171 eggs laid). Loss of eggs,
presumably from snake predation, was higher than loss of
Maguari Stork 95
nestlings from all causes combined. Most nestling losses
were caused by large young falling out of nests that con-
tained 3^ chicks. No brood-reduction through nestling
starvation was observed in any of the 123 nests studied.
Within 3.5-4.5 months after fledging, the young had left the
area (B.T. Thomas 1984), and, although 128 young were
ringed over four seasons, none were seen again in the nest-
ing area until they returned to breed at 3-4 years of age
(B.T. Thomas 1986).
Males apparently become sexually mature at 3 years of
age and females at 4 years (B.T. Thomas 1987).
TAXONOMY
This species shows a number of similiarities to the Oriental
White Stork of northeastern Asia, with which it shares some
similar aspects of morphology and some behaviours, and to
the Woollynecked Stork, with which it shares its distinc-
tively forked tail. The fossil stork Ciconia maltha from the
Pleistocene of North America may provide a past link be-
tween the ancestors of the modern-day Maguari and Orien-
tal White Storks (H. Howard 1942; Brodkorb 1963). Like
the Black Stork, the Maguari apparently clatters only rarely
during the Up-Down display and may not do so in all areas
of its range. As pointed out by C.E. King (1988), these are
the only two members (among those that have been studied)
of the genus Ciconia that habitually nest in heavily vegetated
sites; in such situations it might be advantageous to possess
displays that are less conspicuous to potential predators that
might not otherwise find the nest over a long range.
There has been much confusion regarding the scientific
name of this species. It was formerly called Euxenura galeata,
after Molina (1782). However, this description was of a
composite nature (Hellmayr and Conover 1948, Kahl
1979b) and cannot be used. Following the suggestion of
Delacour and Mayr (1945) and Kahl (1972e) we have
merged the monotypic genus Euxenura into Ciconia, with
which it shows a close relationship. This decision was
further supported by the work of D.S. Wood (1983, 1984)
and B.T. Thomas (1988).
CONSERVATION
Status throughout the range is poorly known, but in the
Venezuelan Llanos, where the population was studied for
13 years, it has decreased markedly within the past decade
(Luthin 1984a, B.T. Thomas 1986, 1987). Aerial censusing
of nesting areas could add greatly to our knowledge of its
overall status. Recently, the first chicks hatched in captivity
in 30 years were produced at Disney World, Florida (Healy
1991).
The Maguari is considered a gamebird in Amazonia (Sick
1985) and elsewhere. Hunting (including the taking of
young for food), human disturbance and loss of habitat to
agricultural activities are primary threats, and possibly the
causes of population decreases seen in recent years (B.T.
Thomas 1986, 1987).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 295.
 White Stork
Ciconia ciconia (Linnaeus)

Subspecies:
Ciconia ciconia ciconia (Linnaeus)
Ardea Ciconia Linnaeus, 1758, Syst. Nat., ed. 10, p. 142: Europe, Asia, Africa: restricted to Sweden
by Linnaeus, 1861, Fauna Svecica, ed. 2, p. 58
Ciconia ciconia asiatica Severtsov
Ciconia alba asiatica Severtsov, 1873, Izvestiia Imp. Obshchestva Liubitelei Estest. Antrop. Etnogr.,
Moscow, 8, pt 2 (1872), p. 113: Turkistan

Other names:
European Stork (English); WeiBstorch, WeiBen Storch, Adebar (German); Gigogne blanche
(French); Ooievaar (Dutch); Ciguena comun (Spanish); Cegonha branca (Portuguese); Ciconia
bianca (Italian); Hvid stork (Danish); Vit stork (Swedish); Bely Aist (Russian); Capa bfly
(Czechoslovak); Bocian bialy (Polish); Roda bijela (Yugoslav); Barza alba (Rumanian); Feher golya
(Hungarian); Pelargos (Greek); Bajbar, Badjah, Belaridj, Haji, El-haz laqlaq, Laklaka, Najeh,
Mehab (Arabic); Leklek, Bu-laqlaq (Turkish); Lag-lag, Haji lag-lag (Hindi); Mashowori (Shona);
Witooievaar, Grootsprinkaanvoel [='Great Locust Bird'] (Afrikaans); Lima (Hausa)

IDENTIFICATION
Standing 100-125 cm tall, this large, mostly white, stork has black flight feathers in the wings, a straight
red bill, and long red legs.
The adult plumage is all white or near-white, except for the primaries, secondaries, greater primary
and secondary coverts, alula, and scapulars, which are black glossed with green or purple. There is
sometimes a small, variable amount of white or silvery grey on the outer vanes of the secondaries and/or
inner primaries. The feathers of the lower foreneck and chest are elongated, forming a ruff that can be
erected.
The bill, legs and feet are red to orange-red; the legs may be whitish, owing to coating with uric acid
from excrement. The iris is brown, and the bare skin surrounding the eye and on the lores is black. The
naked gular skin is black, grading to orange-red posteriorly and along the border adjacent to the
feathered neck.
The sexes are similar except that the male is usually larger and with a more massive bill, which is
sometimes subtly upcurved on the ventral border of the lower mandible (Schierer 1960a). In pairs
where morphological differences are obscure, the sexes can often be distinguished on the basis of
behavioural differences in ritualized displays (Schiiz 1942b, Kahl 1972c).
The subspecies asiatica is on average larger than ciconia and its bill is redder, upcurved, longer and
heavier.
There are no dramatic differences between the non-breeding and breeding plumage, except that in
the latter the whites of the feathers are cleaner and the areas of bare skin are more brightly coloured.
The white plumage, especially the neck-ruff, becomes soiled as the breeding season progresses.
The first-down of hatchlings is short, sparse, and off-white; the second-down (at 1 week) is dense,
woolly, and white. Weight at hatching is 70-77 g (O. Heinroth 1922:186, Dementiev and Gladkov
1951:450, Schropel 1984). By the third week, black scapulars and flight-feathers, begin to appear. At
hatching the bill'is black with a brown tip, and legs are pink, later becoming grey-black.
98 White Stork
Juveniles are similar to adults but the black wing-coverts
and scapulars are tinged with brown, and the primaries and
secondaries are less glossy. The bill is, at first, shorter than
the adult's and changes gradually from blackish to brownish
or pale red with a blackish tip. The legs are a duller
brownish red. Measurements of developing captive young
are given by L. Gangloffet al. (1989).
The only true vocalization by adults is a hissing, given
primarily at the nest in conjunction with some forms of the
Up-Down and other displays. Loud and prolonged clatter-
ing of the bill produces a rapid machine-gun-like rattle, with
a marked crescendo and diminuendo. The tongue-bone
expands the gular pouch, which acts as a resonator for the
sound during clattering. Nestlings give a Begging call remi-
niscent of the mewing of a cat; they also clatter their soft
bills, but without much sound.
White Storks fly with neck and legs fully extended. They
often soar high on thermals of rising warm air, especially
during migration. Between thermals they glide in loose
scattered formation, not usually in rows or Vs as some other
soaring birds. During flapping flight, slow, steady wing-
flaps are interspersed with short glides. The flapping rate
averages 170/min (27 recordings) (Kahl 1971b); the flight
speed has been timed at 40-47 km/h (6 recordings) (Penny-
cuick 1972). Migrants have been reported at 4300 m in the
Himalayas (Meinertzhagen 1955) and at 3000m in the
southern Sudan (J. Vincent 1948). They have also been
seen at 3300 m in the western Sudan (J. Vincent 1948) and
at 1600 m near the Dead Sea, where one flock was mistaken
for enemy planes (apparently during World War I) and
fired on for ten minutes, one stork being killed (Meinertz-
hagen 1954:389). M.I.M. Turner (1964), while flying a
small plane over the Serengeti Plains, Tanzania, encoun-
tered White Storks at 3600 m above the ground. Birds
occasionally clatter during flight, especially in the vicinity of
the nest-site (pers. obs. M.P.K.).
Within their breeding range, White Storks might possibly
be confused with one of the white egrets, which have no
black in the wings. In the winter range, they could be
confused with the Yellowbilled Stork, which has a decurved
yellow bill, naked red face, and a black tail. When soaring
high, they might be confused with the Great White Pelican
(Pelecanus onocrotalus) or the Egyptian Vulture (Neophron perc-
nopterus)) which show similar wing-patterns, but the ex-
tended long neck and legs of the stork set it apart.
DISTRIBUTION AND POPULATION
The overall distribution of this species includes much of
Europe, the Middle East, west-central Asia, northwestern
and extreme southern Africa. Vagrants have occurred in
Iceland, Britain, Ireland, Norway, Finland, Malta, Azores,
Madeira, Canary Islands, Cape Verde Is., St Helena I., and
Sri Lanka.
Ciconia ciconia ciconia is found in Europe in southern and
eastern Portugal, western and central Spain and eastern
France; in The Netherlands, Denmark, eastward to the Gulf
of Finland and in the Pskov/Leningrad regions of the USSR,
southward to the Sea of Azov, Turkey, northern Greece,
eastern Yugoslavia, sporadically occurring in northern
Italy. Formerly it bred in southern Sweden, Switzerland,
western France, Belgium and southern Greece. In north-
western Africa it is found in northwestern Tunisia, northern
Algeria, northern and northwestern Morocco. In the Mid-
dle East it is found in Turkey and Azerbaijan, USSR,
southward to western Iran, and, increasingly, to Israel. A
few pairs breed in the extreme southern tip (Bredasdorp,
Mossel Bay and Oudtshoorn regions) of Cape Province,
South Africa. The reports of breeding in Zambia, Zim-
babwe and Zululand, South Africa, reported in Mackworth-
Praed and Grant (1962:79) were in error (Clancey 1964b).
It may have bred in Kenya in 1982 (Dittami and Haas
1982). A few birds winter in the southern parts of the
breeding range; most migrate to tropical Africa or, in lesser
numbers, to southwestern Asia.
Ciconia ciconia asiatica is found in Turkistan, USSR, Uzbek
SSR, Tadzhik SSR, Kirgiz SSR, and southern Kazakh SSR,
from Amu Dar'ya to Issyk KuF and Lake Balkhash and
extreme western Sinkiang, China (Kashgar). The summer
range covers approximately between 39^3°N and 59-79°E
(Creutz 1988:17). It migrates through Afghanistan and win-
ters mainly in the northern part of the Indian subcontinent,
mixing with birds of the nominate race from the Middle
East. As many as 360 White Storks (race not determined)
were seen in extreme southern India in January 1981 (J.M.
Johnson 1984). Two White Storks with red bills seen west of
Bangkok, Thailand, in January 1964, may have been from
this population (P. Round et al. unpub. report).
The White Stork is a long-range, diurnal migrant, which
is very much dependent on soaring/gliding flight, assisted
by rising thermals of warm air, for covering long distances
with relatively little expenditure of energy. Movements take

place mainly during the warmer times of the day, and birds
can transit 100-200 km per day. Probably owing to this
reliance on thermals, most European birds pass around
both ends of the Mediterranean; only a few cross in the
middle (see map in Schulz 1988:16). When water must be
crossed, storks generally do so at the narrowest point.
The main departure from the European breeding grounds
is in August; as a rule juveniles leave before adults. A
majority of birds from eastern Europe cross the Bosphorus,
go around the eastern end of the Mediterranean, and enter
Africa across the southern end of the Gulf of Suez. Whereas
most of those from extreme western and southwestern Eur-
ope cross at the Straits of Gibraltar. The migration-divide
(German: Zugscheide) in Europe is discussed in detail in
Schuz(1963a).
Autumn migration at the Bosphorus peaks between mid-
August and early-September. It is estimated that approxi-
mately 370000 adult and immature White Storks pass
through this area each autumn (Goriup and Schulz 1990).
Flocks of up to 11 000 storks and total numbers of over
207 000 were observed between 23 July and 6 October 1966;
80% of the movement took place between 09.00 and 13.00 h
(Porter and Willis 1968). Up to 315000 were counted at the
Bosphorus in the autumn of 1972 (Kasparek and Kilic
1989a). At Burgas Bay, northwest of the Bosphorus in
Bulgaria, 226956 were counted between 10 August and 30
October 1981; during the 5 years 1979-83, peak migration
dates fell between 21 and 31 August; flocks averaged 577,
but one flock contained 21 000 birds (Michev and Profirov
1989). On the western route, 27414 birds were counted
crossing the Straits of Gibraltar between 20 July and 4
September 1985 (Lazaro and Fernandez-Cruz 1989).
Birds using the eastern route winter mainly in the eastern
half of Africa, many as far south as Cape Province, South
Africa. In extreme cases, the round-trip journey from Euro-
pean breeding grounds to wintering areas in southern Africa
may be over 22 000 km. It was estimated that a 25-year-old
bird from East Germany may have covered over 500 000 km
on migration in its lifetime (Creutz 1979b).
Birds using the western route winter mainly in the north-
ern tropics (12-17°N) of West Africa (Moreau 1972:240;
ringing recoveries summarized in Cramp 1977:330, and
Kanyamibwa et al. 1990). The return journey in spring is
more rapid, with most birds arriving back on European
breeding grounds in March-April. In certain years adults
return late to the breeding sites and their nesting success is
seriously reduced; the cause of these 'disturbance years'
(German: Storungsjahre) is not known but may relate to
unfavourable conditions on the wintering grounds or during
migration.
Wintering birds in Africa move around to a large extent
in response to changing abundance of food, such as locust
swarms (Schiiz 1959a,b). When food is abundant, the birds
may gather in very large numbers: for example, 100000
White Storks were estimated in a 25 km2 area of northern
Tanzania in January 1987, feeding on an army-worm infes-
tation (Schulz 1988:17, 364, D.A. Turner pers. comm.
1990). There is some evidence that young birds often mi-
grate further south than adults (Schiiz 1959a), and non-
White Stork 99
breeding immatures often remain in the winter range during
the northern summer. One juvenile, ringed as a nestling in
extreme southern South Africa, was recovered 4 months
later near the Tanzania/Zambia border, approximately
3100 km to the north (McLachlan 1963).
Extensive ringing in Europe over many decades has fur-
nished us with valuable information on movements, longev-
ity and life-history. However, recent reports indicate that
ringing may result in increased mortality because the rings
can injure the bird's legs (Schulz 1987, Lohmer 1989). It is
also possible that some birds are shot, out of curiosity,
specifically because they are wearing a ring (D. Jonkers
pers. comm. 1989).
Counts of breeding birds in Europe are extensive and
often of long duration. International Stork Censuses have
been conducted in 1934, 1958, 1974, 1984 (Schiiz 1936a,
1979b, Schiiz and Szijj 1960a, Boettcher-Streim and Schiiz
1989, Rejman and Folk 1991). The trend is generally down-
ward in northwestern Europe, with the last regular breeding
in Belgium in 1895, Switzerland 1949, and Sweden 1954.
Populations appear to be declining at a slower rate in
eastern Europe (they may even be increasing and extending
their range to the north and east in the Baltic states and
European USSR).
In The Netherlands numbers dropped from about 500
breeding pairs in 1910 to 209 in 1929, 85 in 1950, and 5 in
1985 (Jonkers 1989). In Alsace, France, the population
declined from 177 pairs in 1947 and 145 pairs in 1960, to less
than 20 pairs in 1986 (Schierer 1986, Kanyamibwa et al.
1990). In western Germany the number of breeding pairs
dropped from about 2500 in 1958 to only about 500 in 1987
(Schulz 1988). And in Denmark numbers went from about
4000 breeding pairs in 1890, 1000 in 1930, 222 in 1952, 111
in 1964, 54 in 1971, 19 in 1984, to 12 in 1989 (Schiiz 1986c,
H. Skov pers. comm. 1990).
Numbers have increased in some areas of eastern Europe.
In Estonia, the first nest was found in 1841; there were
approximately 320 pairs in 1939 and 1954, and by 1974
there were 1060 pairs. Similarly, in the Pskov Region of the
RSFSR, numbers climbed from 400 pairs in 1958 to 1218
pairs by 1974 (Veroman 1976).
Population figures through the mid-1980s are summar-
ized in Rheinwald et al. (1989). These recent estimates are
(breeding pairs): Spain 6753, Portugal 2004, France 19, The
Netherlands 2, Denmark 14, western Germany 649, eastern
Germany 2722, Austria 318, Poland 30500, Czechoslovakia
1670, Bulgaria 5422, Rumania 3700, Hungary 4693, Yugo-
slavia 4500, Greece 1500, USSR 37086, Turkey 9000, Iran
2394, northwest Africa 15830. The total from these surveys
is approximately 128000 breeding pairs. Some areas of the
USSR and Middle East were not included, because of lack
of data. About 19000 of these pairs belong to the western
European population (birds nesting west of the Zugscheide),
migrating through Gibraltar to West Africa. Between 1974
and 1984 this western population declined by 20% on aver-
age, considerably more in the north than the south. The
eastern population, which migrates via the Bosphorus, de-
clined by about 12%, more in the south than the north
(Rheinwald 1989b).
100 White Stork
At the fringes of breeding range, there are about 6 pairs in
the South African breeding population (Siegfried et al.
1976) and an estimated 600+ pairs in the C. ciconia asiatica
population (Schiiz and Veromann 1982).
Wintering numbers are more poorly known. Estimates of
numbers wintering in Africa are at least 450 000 individuals
in the eastern population and 90000-100000 in the western
(Schulz 1988). A total of 310 000 White Storks were counted
passing through Israel in the spring of 1984 (Paz 1987:33),
with some remaining throughout the year (pers. obs.
J.A.H.). Between 20000 and 24000 birds winter in the
Kirov Bays area of the southwestern Caspian sea, about
140 km south of Baku (Goriup and Schulz 1990).
According to a recent survey .{Goriup and Schulz 1990),
the White Stork population level today is under half of what
it was 40 years ago and is still falling.
ECOLOGY
The White Stork favours open country. In breeding areas, it
is most often found in wet pastures, moist meadows, shallow
lakes and marshes; it is fond of fields during ploughing or
mowing. Feeding areas are usually adjoining or close to the
nesting site. In the winter range, it is more often found in
drier savannas and open grasslands, sometimes gathering at
lakes, ponds or rivers.
Food locating is mainly visual. A bird Walks Slowly or
rapidly forward, with measured strides, holding its head
forward of the body and the bill pointed down. Sometimes
the White Stork locates aquatic prey by tactolocation, in a
manner similar to the wood storks (Mycteria) (Lohrl 1957).
Foraging is done singly, in small flocks, or in large groups if
prey is abundant; at locust swarms up to 2000 have been
reported in Uganda (FJ. Jackson 1938:69), 5386 in South
Africa (Milstein 1966), and tens of thousands in northern
Tanzania in January (Moreau and Sclater 1938). It is
normally a diurnal feeder but is also active at night, by
moonlight, during the breeding season (Darazsi 1958, Golo-
dushko quoted in Pinowska and Pinowski 1989, Bauer and
Glutz von Blotzheim 1966:405). Foraging efficiency of juv-
enile storks in Poland was found to be significantly lower
than that of adults (Pinowska and Pinowski 1989).
The White Stork's food is small animals, the diet varying
considerably with feeding conditions. Prey includes insects
and their larvae (especially beetles, grasshoppers, locusts),
reptiles (lizards, snakes), small mammals (especially voles,
mice, moles, shrews, young rats), and earthworms (espe-
cially important in early spring and during flooding). Less
commonly it eats the young and eggs of surface-nesting
birds, frogs (an important food item in some areas, not in
others), toads, tadpoles, fish, molluscs and crustaceans.
Indigestible remains of food are later regurgitated as pellets,
measuring about 50 X 34 mm and weighing about 16 g
(Haverschmidt 1949:47).
Grasshoppers and locusts seem to be the most important
food items on the African wintering grounds. Two storks
examined had 1000 and 1600 locusts in their stomachs
(Mell 1951). A first-year bird in western Kenya had eaten so
many locusts that it was unable to fly and was killed by a
'wild cat' (Schiiz and Bohringer 1950). There is some evi-
dence that storks may store excess fat before migrating.
Milstein (1966) found birds that had been feeding on abun-
dant locusts to be very fat in February in South Africa.
Adults in Europe seem to gain weight late in the breeding
season: in the month of June, 14 males weighed an average
of 3.341 kg, 14 females 3.15kg; in July, 12 males averaged
3.97 kg and 12 females 3.521 kg (G. Niethammer 1938:291).
Birds are often very gregarious on migration and in the
winter quarters. They sometimes gather in large numbers,
hundreds or even thousands, at abundant food sources such
as locust swarms and grass-fires. The stork rests, often on
one leg, with the bill buried in the ruff of feathers on the
lower foreneck; both of these postures probably help to
minimize heat loss at low ambient temperatures (pers. obs.
M.P.K.). They are able to swim although rarely do so
(Zimmermann 1955).
Habitat losses in European breeding areas are probably
very important in causing population declines. However, an
analysis of long-term population and breeding trends in
northwestern Europe suggests that 'critical mortality occurs
in the winter in Africa' (Lack 1966:228). Changes in African
feeding conditions, owing to factors such as droughts and
locust-control, may be of major importance (Dallinga and
Schoenmakers 1989).
On migration, particularly in Europe, collision with wires
and electrocution are important causes of mortality (K.
Brouwer and C.E. King pers. comm. 1990). Hailstorms
sometimes cause mass mortality of wintering birds in South
Africa (J. Wittmann 1932, Badenhorst 1956, Kuhk, 1956).
The heat and adverse winds (Arabic: khamsins] of desert
crossings also take their toll of migrants (Schiiz 1935b,
Bannerman 1957:11-12, C.A. Reed and T.E. Lovejoy
1969:152). Particularly on migration, there is evidence that
storks are sometimes attacked by birds of prey (Schiiz
1965a, 1975, Breach 1970, Harwin 1970), and many are also
killed by humans (Hartert 1912-3, Giraudoux and Schiiz
1978, Schulz 1988). A total of 484 ringing recoveries from
southern Africa shows mortality rates in birds which died
there to be 59% in the first year, 47% in the second, 36% in
the third, and 20% in the fourth to tenth years, after which
few recoveries were made (Oatley 1989). Other analyses
indicate a first-year mortality between 55% and 74%
(Schiiz 1955a).
BREEDING
Usually the White Stork nests in the lowlands, but it does
nest at up to 2000 m in Morocco (Heim de Balsac and
Mayaud 1962:74) and Armenia, USSR (Dementiev and
Gladkov 1951:446). Especially in Europe, the White Stork
often breeds in close association with man, on rooftops,
towers, chimneys, telephone-poles, high-wire towers, walls,
hay-stacks and special poles erected for the purpose. In
Armenia in 1988, 53% of nests were on poles and other
telegraph wire supports, 29% on traditional buildings and
18% on modern structures (Adamian 1991). It also some-
times nests on trees, atop cactus (e.g. in Spain), or on cliff

ledges, and, very rarely, on the ground (Thienemann 1911).
Nests are built singly or in small, loose colonies; in Spain as
many as 9 nests have been found on one rooftop (Chozas,
1984), and colonies may contain up to 30 pairs (Goriup and
Schulz 1990). Some nests have been in use annually for
hundreds of years; one known to exist in 1549 was still
occupied in 1930 (Goriup and Schulz 1990). The bulky nest
is constructed of branches and sticks, lined with twigs,
grass, paper and rags; it is reused in successive seasons and
added to each year and throughout the nesting season. Nest
platforms are approximately 0.75-1.7m (exceptionally to
2.25 m) in diameter and 0.5-2.0 m (exceptionally to 2.8 m)
high; the size increases with age. One nest, taken down from
a cathedral weighed 800kg (Madon 1935). European nests
are known to have been in use regularly for over 100 years
(Naumann 1838). Ringing recoveries in central Europe
show that about 80% of first-breeders settle on a nest site
within 50 km of where they were hatched (Hornberger
1943a, Schiiz 1949c). Nesting is sometimes in mixed colon-
ies with other species, such as with Cattle Egrets (Bubulcus
ibis] in Spain.
Pair-bonds are monogamous; rarely extra females attend
a nest (Thienemann 1911, Assfalg and Schiiz 1988).
Partners apparently do not migrate or winter together but
often re-mate in successive years, probably owing to mutual
attachment to the same nest-site. Males usually arrive back
at the nest-site first. Sexual maturity is usually attained at
3-5 years; one male bird was still breeding successfully at
25 years (Creutz 1979b).
Nesting (and sometimes roosting) birds greet newcomers
with loud bill-clattering and Up-Down displays. Strange
storks that approach an occupied nest often trigger a violent
attack by its owner. If a newly-arrived female is quickly
accepted by a male at a nest, it may signify that she is a
returning mate from a previous season. Unwelcome
strangers are threatened with a version of the Up-Down
display in which the wings are held away from the sides and,
especially in the male, pumped rhythmically up and down.
Thus, the Up-Down display, in its various forms and
contexts, can function as an advertising display, a threat or
a greeting between mates (Camanni et al. 1989). When used
as a greeting between mates, the Up-Down probably func-
tions to suppress aggression (pers. obs. M.P.K.). The
throat-sac is extended by the tongue-bone and acts as a
resonator during clattering (Siewert 1932b:136).
When a strange female attempts to approach an unmated
male on a nest, he often displays by giving the Head-
Shaking Crouch, orientated toward the female. In this dis-
play, the male drops to the nest, as if incubating, fully erects
the feathers of the neck-ruff, and shakes his head vigorously
with a side-to-side motion. If, eventually, the female is
accepted by the male, mutual Up-Down displays follow and
soon thereafter copulations. A successful copulation lasts
about 20 s (Schiiz 1934), and there may be as many as 107
day (Schiiz 1942b). Copulations have been witnessed, in
wild pairs, with the female standing, sitting on her tarsi or
lying flat on the nest (pers. obs. M.P.K.). After a pair is
formed, mutual allopreening, especially in the feathers of
the head and neck, is common between mates. For further
White Stork 101
details on ritualized displays observed during courtship, the
reader is referred to Schiiz (1942b) and Kahl (1972c).
Nest-building is by both sexes; males bring most of the
nest-material (Nagy 1955). Although old nests are usually
reused, a new nest can be constructed in as little as 8 days
(Goriup and Schulz 1990).
A clutch contains 3-5 eggs, usually 4, the extremes being
1-7. Eggs are chalky white; they are laid at intervals of 1-4
days, usually 2 days. Of 80 clutches in The Netherlands, 5%
had 2 eggs, 25% 3, 48% 4, 16% 5, 6% 6; the average was
3.93 eggs per clutch (Haverschmidt 1949:39). Of 837
clutches in southwestern Poland, the figures were: 1% 2,
48% 3, 44% 4, 8% 5; the average was 3.58 eggs per clutch
(Mrugasiewicz 1972).
The incubation period usually lasts 31-34 days. In Spain,
97 eggs took between 24 and 33 days to hatch, and the
average was 30.4 days (Chozas 1986a). Incubation is by
both sexes, female probably at night; it starts with the first
or second eggs, and hatching is asynchronous. Young bill-
clatter to parents from day one (Siewert 1932b:155). Both
parents feed nestlings by regurgitation onto floor of nest.
Nestlings are fed about once per hour when 5-6 days old
and about once every 2 h when 15-16 days old (Schiiz
1943c). Water is regurgitated to the young on hot days
(Schiiz 1943c, Haverschmidt 1949:50, Dementiev and Glad-
kov 1951:450). Parents often forage and feed their young on
clear nights (Schiiz and Schiiz 1932b). Young are guarded
constantly by one adult during their first 2-3 weeks. The
fledging period is 54-70 days; young become independent
and leave the area within another 7-20 days (G. Nietham-
mer 1938, Dementiev and Gladkov 1951, Bauer and Glutz
von Blotzheim 1966, Cramp 1977). Additional details on the
care and development of nestlings may be found in Schiiz
(1943c). The elapsed time from the completion of the clutch
until first flight of the young averaged 91-92 days, based on
540 records (A. Mrugasiewicz pers. comm. 1969).
There is considerable mortality (mainly of eggs, but also
of some nestlings and even of adults) during violent fights
between owners and strange storks that often attack and
attempt to take over occupied nests (Schiiz 1936b). In East
Prussia during 1931, 2660 cases of fighting were recorded at
8835 nests, with 77 adult birds left dead (Schiiz 1944).
These fights are strongest between adult storks of the same
sex (Schiiz 1944).
Nesting success is higher in years when birds return
earlier to their breeding sites from the winter quarters. In
nests with several young, the smallest is reared successfully
only in years when food is plentiful; when all young in a nest
die, it is often the result of infestation with intestinal trema-
todes from frogs in the diet (Szidat 1935, Schiiz 1936b;
additional discussion of parasites is given in Szidat 1940,
1943). When environmental conditions are optimal, as
many as 6 young may be fledged from a single nest
(Schneider 1988 cited in Goriup and Schulz 1990). In Eur-
ope overall, the number of young reared varied from 0.8 to
3.4 and averaged about 2 (Bauer and Glutz von Blotzheim
1966:404) There are several long-term studies of nesting
success. In Germany (Oldenburg) during 1928-63, the
brood size reared varied from 0.8 to 2.84, with a mean of
102 White Stork
1.94 (Tantzen in Lack 1966:216, Schiiz 1981a:86); in The
Netherlands during 1934-45, from 1.3 to 2.6, mean 1.96
(Haverschmidt 1949:52) and during 1950-75, from 1.1 to
2.9, mean 1.7 (Jonkers 1975:265); in Denmark during 1952-
71, from 1.3 to 2.9, mean 1.9 (Dybbro quoted in Cramp
1977:335). Breeding success appears to be highest when the
parents are 10-16 years old (Meybohm and Dahms
1975:60).
TAXONOMY
The Oriental White Stork, in the Far East, has often been
treated as a subspecies (C. ciconia boyciana) of the European
bird. However, recent studies (e.g. Archibald and Schmitt
pers. comm. 1990, G.E. King 1988) have shown the mor-
phological and behavioural differences to be so great that we
feel it should be considered a full species (Ciconia boyciana).
Existing data suggest that the storks of the Central Asian
population are considerably larger (see Appendix) than the
European population (Vaurie 1965:84); some authors (e.g.
Meinertzhagen 1930:432; J. Niethammer 1967b) doubt
whether the differences are sufficient for distinction. Tenta-
tively, we do recognize it as a subspecies (asiatica); however,
additional measurements and other information on the
bird's biology are desirable.
The genus Ciconia as now constituted includes storks that
have, at times in the recent past, also been classified under
the genera of Sphenorhynchus (Abdim's), Dissoura (Woolly-
necked and Storm's), and Euxenura (Maguari) (Delacour
and Mayr 1945, Kahl 1972Q.
CONSERVATION
The biology of the White Stork is one of the best known of
all birds (Zink 1967, Rheinwald et al. 1989:470), primarily
because of the species' close association with man, and its
large size, popularity and abundance in Europe. In spite of
this extensive scientific knowledge about the bird, we have
not been able to prevent a continued decline in numbers
over much of its range. Essentially it has been extirpated
from Sweden, Denmark, The Netherlands, France, Switzer-
land, Belgium; and, if present trends continue, it may be
gone from western Germany by the end of this century (C.
Luthin pers. comm. 1989).
Reasons for the decline in Europe are not firmly estab-
lished (see discussions in Dallinga and Schoenmakers 1987,
1989, Goriup and Schulz 1990), and it is probable that
several independent causes are involved. The following fac-
tors have been cited by various authors: destruction or
alteration of feeding areas in breeding range; reduced food
supplies in winter range (owing to droughts and insecti-
cides); chemical pollution, in both summer and winter
ranges; electrocution and collision with overhead wires;
shooting (mainly on migration). It would seem most likely
that it is loss of habitat—owing to development, industrial-
ization, and intensive agriculture—in western Europe that
has caused much of the decline (C. Luthin pers. comm.
1989). Otherwise why would there be such a distinct differ-
ence in numbers of storks at the borders of eastern and
western Europe? It remains to be seen what effects the
recent political changes in Europe will have on stork popu-
lations. Storks will undoubtedly suffer if future economic
growth in eastern Europe is not tempered with ecological
sense.
Whatever the reasons for the decline, it would appear to
be largely caused by man. However, long-term climatic
changes may also be involved, especially in the protracted
decrease of populations wintering in the West African Sahel,
where there has been a generally declining trend in rainfall
during 1943-85 (Kanyamibwa et al. 1990). Lack (1966)
analysed data from German stork populations and con-
cluded that the reasons for the decline should be looked for
in Africa. Goriup and Schulz (1990) summarized it best
when they said that 'western birds suffer most since they
find no respite in either western Africa or the western
European breeding grounds, whereas the eastern birds have
the opportunity to compensate for poor breeding success by
better survival in eastern and southern Africa during the
winter'.
In many areas of Europe and the Middle East, the White
Stork is revered as a symbol of good luck and prosperity and
is protected from direct persecution by popular custom.
People sometimes say of an unlucky man: 'he must have
killed a stork', and in ancient times the penalty for killing
one was the loss of a hand (Gary 1973:38). Unfortunately
the killing of a stork carries no such stigma in many parts of
the world where the birds go during their non-breeding
seasons, and many are killed by humans (Schulz 1988).
However, direct killing by humans is of slight significance in
the overall mortality of White Storks (Goriup and Schulz
1990).
Attempts are being made to re-establish artificially the
White Stork as a breeding bird in some areas of Europe,
primarily Switzerland, France, The Netherlands and Italy,
where it has disappeared (see Schierer 1972, Bloesch 1980,
1989a, Vos 1989, Camanni and Tallone 1990b, P. Arnold
1990, Blackwell 1990). Perhaps the most successful results
have been achieved by Max Bloesch, of Switzerland, who
re-introduced young birds taken from nests in Algeria, and
—as last reported (1985) —had 108 nesting pairs of free-
flying birds (Bloesch 1989a). Most of these birds remain the
entire year in the immediate area and are artificially fed, so
they cannot be equated with a truly 'wild population' that
migrates to Africa each winter (cf Rheinwald, 1989a:436).
Since the offspring of these birds sometimes do migrate
away in the winter, it remains to be seen if a truly wild,
naturally behaving population will eventually become re-
established. This is, of course, the hope.
Such captive reintroduction programmes are contro-
versial with many biologists. Some doubt whether 'project'
storks can ever become truly independent of care by man.
There are concerns about the wisdom of introducing foreign
gene pools (e.g. Algerian storks into Switzerland). Since
they often winter in the area and occupy nest-sites early in
the spring, 'project' storks may out-compete wild migrants
for nesting areas in regions where these are in short supply.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
pages 296-297.
 Oriental
White Stork
Ciconia boyciana Swinhoe

Ciconia boyciana Swinhoe, 1873, Proc. Zool. Soc. London, p. 513: Yokohama, Japan

Other names:
Eastern White Stork, Japanese White Stork, Japanese Stork, Chinese White Stork, Korean White
Stork, Blackbilled White Stork, Blackballed Stork, Far Eastern Stork (English); Schwarzschnabel-
storch (German); Kitaisky Bely Aist, Dalnevostochny Bely Aist (Russian); Kono-tori, K6
(Japanese); Pai-kuan, Bai-guan (Chinese)

IDENTIFICATION
Standing 110-150 cm tall, the Oriental White Stork is a very large white bird with black flight feathers
in the wing. The bill is black and the legs and feet dark red.
Adult plumage, including tail, is white or off-white; the primaries, secondaries and greater wing-
coverts are black with a slight purplish or greenish sheen. Several of the inner primaries and the
secondaries have the upper side of their outer vanes silvery grey to white (more pronounced than in the
White Stork). At the base of the foreneck, there is a loose ruff of longer feathers, often soiled in
appearance, that are sometimes erected during displays. The bill is black, tinged purplish or deep red at
the base, slightly upcurved and, sometimes, with a slight gap between the mandibles in the middle
(Vaurie 1965:85). The bare skin around the eyes is red and has a pebbly texture. Although there is some
overlap in size, males are usually noticeably larger than females and have a relatively more massive bill.
In the breeding season, there are no major changes in appearance, except that the newly moulted
plumage is cleaner and the soft parts become more brightly coloured. The naked skin around the eye,
lores, and on the throat-pouch becomes a deeper, richer red. The legs and feet are dark red. The iris is
pale yellow-grey to pale blue.
At 1 day of age, the nestling is covered with dirty white down and weighs 82-88 g. The bare skin near
the eyes is a light pinkish orange and the iris is dark brown. The bill at first is ivory to greyish white with
a yellow hue and later becomes a dirty greenish and yellow-orange in colour. The legs and feet are
yellowish pink at first and later become light orange. By 16 days the nestling is covered with pure white
down, and the bare parts of the head are a bright reddish orange. By 62 days the bill has become a dark
grey with a yellowish brown dorsal edge, and the iris is a light yellowish grey. Further details on the
growth and development of the nestlings are given in Winter (1982, 1991). C.E. King (pers. comm.)
found that at fledging, captive young had plumage similar to that of adults, but they had brown on some
feathers; their legs, feet, bill and bare areas of the face did not achieve the full colour of adults. Juveniles
are described as being similar to adults, except that black feathers show a brownish cast and there are
brown patches on the scapulars (Dementiev and Gladkov 1951:455).
A loud and prolonged clattering of the bill is heard during many displays at the nest; clattering is also
heard, occasionally, at feeding areas or in flight. By lowering the hyoid apparatus into the base of the
gular region, a sound chamber is created that gives the clattering a reverberative component (C.E. King

Shaded red area indicates former range.
1988:53). A 'hissing* vocalization is a component of agonis-
tic displays by adults in captivity (C. King pers. comm.
1990, K. Archibald pers. comm. 1990).
A *Kaa, kaa' or £Yia, yia' Begging call is heard from
young, starting several hours before they hatch. A few days
later a hoarse 'Yeee' call is given, growing louder and lower-
pitched with age (Winter 1982, Fei et al. 1991b). C.E. King
(1988:105) described the Begging call of captive nestlings as
'a bisyllabic vocalization that resembled a cat's "meow" in
cadence1.
In migration, Oriental White Storks fly in a scattered
formation, unlike the V-formations of cranes (Hemmingsen
195la). In China, birds within the same flock soaring to
gain height in thermals circle in both clockwise and counter-
clockwise directions. Migrants progress with both flapping
and gliding flight. Flocks of autumn migrants often con-
tained several hundred birds (M.D. Williams et al. 1991).
On the wintering grounds, they sometimes clatter their bills
when gliding (Wang 1991).
This species might be confused with the white cranes
(Grus japonensis or G. leucogeranus), with which it sometimes
migrates; however, the cranes have less black in the open
wings, and they usually call loudly in flight and form V-
formations, whereas the storks do neither (Hemmingsen
and Guildal 1968:188). In southwestern parts of the winter
range, the Oriental White Stork may mingle, occasionally,
with the White Stork. The Oriental species is considerably
larger and has a black (as opposed to a red) bill. It is not
easily confused with the Milky Stork or Asian Openbill
Oriental White Stork 105
Stork, which occur in the extreme southern portion of the
Oriental White Stork's winter range.
DISTRIBUTION AND POPULATION
The Oriental White Stork breeds in the Amur and Ussuri
River regions of the Soviet Far East (westward to Bla-
goveshchensk and southward to Lake Khanka) and extreme
northeastern China (Heilongjiang, Jilin Provinces), and for-
merly bred in Korea and Japan. It winters mainly in the
Changjiang (=Yangtze) valley, China, a few wintering
south to Fukien Province and rarely to Hong Kong,
occasionally to Taiwan, southern Ryukyu Is. It has wan-
dered northward to southern Yakutsk and Sakhalin Is.,
USSR, and southwestward to Bangladesh, northeastern
India and southern Tibet (Vaurie, 1965:84—85, Fisher et al.
1969:183, fitchecopar and Hue 1978:73, Neufeldt and Wun-
derlich 1982).
Soviet birds depart from their nesting area in the Middle
Amur River region between late August and mid-
September; other migrants, from further north, move
through the area in early-October (Winter 1982:82). In
Heilongjiang Province, China, spring migrants have been
observed to appear in the breeding area in late-March and
autumn migrants had departed by late-October (Fei et al.
199la). Further south, in Liaoning Province, spring mi-
grants pass rather quickly in March and autumn migrants
pass from mid-September to mid-November; the autumn
migration has more birds and is more protracted (Sun
1991).
Most of the migrants seen passing Beidaihe, in eastern
China, did so between 11.00 and 15.30 h. The autumn stork
migrations there exhibited a wavelike pattern, probably
related to weather conditions, with 71% of the 1986 mi-
gration (1945 out of 2729 birds) passing on only 4 days: 29
October; 3, 6, and 14 November (M.D. Williams et al.
1991).
Most Oriental White Storks apparently winter in the
extensive wetlands near the middle and lower Changjiang
River in south-central China, with a few wandering south as
far as Hong Kong (Chalmers 1986:43). They start appear-
ing in mid-October, and the majority of birds have arrived
by December; they leave for the north in early March
(Luthin 1987, Wang 1991). Occasionally individuals may
winter as far north as Korea (Austin 1948:45, P. Won
1973:17, Sonobe and Izawa 1987:42), southern Liaoning
Province, China (Sun 1991), or Maritime Territory, USSR
(L. Taczanowski 1891-93:974, Dementiev and Gladkov
1951:454). Some birds winter at the mouth of the Ussuri
River, USSR, fishing through the ice (Wilder and Hubbard
1938:422).
In eastern China, at Beidaihe, most spring birds are seen
passing during the latter half of March. Many more storks
are seen passing south in the autumn than are seen going
north in the spring (M.D. Williams et al. 1991). This could
reflect some losses during the winter, but probably means
that the main spring migration occurs elsewhere.
In the days when Japan held a breeding population, those
birds were mainly residents or short-distance migrants,
106 Oriental White Stork
spending the entire year in Japan (Jahn 1942, Haversch-
midt 1949:70, Schiiz 1963b, Neufeldt and Wunderlich
1982).
Populations are small. Once common in Japan, the
species is now extinct there as a breeding bird. The major
decline in the Japanese population took place between 1868
and 1895. Illegal shooting and, later, poisoning by mercury
and other agricultural chemicals were the main causes for
the decline. The last successful breeding in Japan took place
in 1959 (W.B. King 1981). Only 7 birds still remained in the
wild in Japan in 1966 (Sakamoto 1966), and only one in
1970 (Kuroda 1970); it now occurs only as a rare migrant or
winter visitor.
The species was still 'locally common' in Korea through
the 1940s (Austin 1948:46). However, it had disappeared as
a breeding bird in South Korea by 1971, when the male of
the last remaining pair was shot (P.O. Won 1971, Luthin
1987). By some unknown date prior to 1989, it also ceased
breeding in North Korea (K. Sonobe pers. comm. 1989).
In 1978, the USSR population, in the Far East, was
estimated at approximately 660 breeding pairs and was
thought to be declining (Flint 1978). On the USSR side of
the Middle Amur River in the Bureya-Arkhara depression,
Winter (1982:77) found 31 pairs nesting in a 162 km2 area in
1975, a density of about 1 pair in every 5.2 km 2 .
Most of the known nesting birds are on the USSR side of
the border (Neufeldt and Wunderlich 1982); little is known
about the breeding populations in Chinese territory.
Martin Williams (pers. comm. 1989, M.D. Williams et al.
1991) reported a total of 2729 Oriental White Storks migrat-
ing past Beidaihe, Hebei Province, in eastern China during
the autumn (11 Oct-16 Nov) of 1986. This is more than
double the previous estimates of total world population and
may represent most of the birds alive at that time. The
current estimate for world population is about 3000 birds
(Luthin 1987).
A total of 1367 Oriental White Storks was counted on the
wintering grounds, mostly in the Changjiang River area of
China, during January 1989 (D.A. Scott and P.M. Rose
1989:80) and 121 in Hong Kong in 1991 (Chan 1991).
ECOLOGY
The favoured habitats of this species are river valleys, wet
meadows and marshes with scattered clumps of trees. In the
Soviet Far East, they have been observed breeding primar-
ily in river lowlands on islands of forest, in areas with
scattered moss-sedge-cottongrass bogs (Winter 1982). Dur-
ing times of former breeding in Japan and Korea, they
frequented cultivated fields, especially rice-paddies, with
nearby wooded hillsides (W.B. King 1981). Wintering birds
in China are found at lakes, marshes, and, sometimes, sandy
beaches and intertidal areas (Shi 1991).
In the wet spring of 1975, adults in the Soviet Far East
foraged within 100-600 m of their nests; later in the sum-
mer, when conditions were drier, they foraged 3-4 km away.
In the drier spring of 1976, foraging was done 3-7 km from
the nests (Winter 1982).
Foraging birds Walk Slowly in shallow or fairly deep
water, often up to their belly feathers, and make swift
downward and forward pecking movements (Winter 1982).
Food finding is mostly visual (Fei et al. 199la). Sometimes,
especially in water where visibility may be limited, feeding
is apparently by tactolocation. Birds plunged the partially
open bill into the water, as they walked, much in the
manner typical of the Blacknecked and Saddlebill Storks.
Up to 17 plunges per min have been counted, when storks
caught an average of 1.22 ± 1.72 S.D. prey items per 5 min
observation period (n = 9). Handling-time varied from 3-5 s
for a carp to 2-3 min for catfish (Coulter 1990c, Coulter et
al. 1991).
Shi (1991) found that wintering birds foraged mainly in
the early morning (07.00-09.00 h) and late afternoon
(15.00-17.00 h) and rested at midday. One bird, watched
for nearly 10 h, caught 486 food items in 742 attempts, a
65.5% success rate.
The food is primarily fish, especially loaches (Misgurnus
fossilis and sleepers (Perccottus glehni), frogs, aquatic beetles
and other insects, also small rodents, snails, arachnids,
snakes, crustaceans, clams, worms and young songbirds
(Austin 1948:46, Austin and Kuroda 1953:333, Neufeldt
and Wunderlich 1982, Winter 1982, Wang 1991, Shi 1991,
Winter 1991). According to S.W. Winter (pers. comm.
1990) the Oriental White Stork is a specialist, with fish
making up 70-90% of the total number of food items and
80-90% by volume. He has recently compiled an analysis
(Winter 1991) of 2600 food items eaten by storks in the
Amur region, USSR.
In the Soviet Far East, vegetable matter (e.g. leaves,
moss, sticks, seeds) made up a large percentage (70-90%) of
the volume in stomachs of some adults and nestlings (Win-
ter 1982:94). However, most of this material is probably
ingested accidentally during the capture of animal prey,
and, being less digestible, it remains in the stomach for a
long time (S.W. Winter pers. comm. 1990). Wintering birds
in China have been observed purposefully eating plant
material, sand, and small stones (Shi 1991).
A Chinese bird was seen catching a catfish weighing
about 750 g (Fei et al. 199 la). Estimates of daily food-intake
of wild, wintering birds in China ranged from 1.5-3.Okg; c.
70% (by weight) of the food taken was fish (Shi 1991).
Apparently the Oriental White Stork is more dependent
on wetlands and more tolerant of cold than is the White
Stork (Hemmingsen 195la). It migrates south in the
autumn later than the European species and winters further
north; its larger size may be an adaptation to cold weather
(M. Williams pers. comm. 1990). Wintering birds, in south-
central China, roost in flocks in open shallow water (Wang
1991).
BREEDING
In the Soviet Far East, nests are placed atop lone trees or at
the borders of woodlands; they have not been closer than 4.5-
5.5km to human settlements (Winter 1982). On the other
hand, in Korea, nests were sometimes built near houses or
villages (Austin, 1948:45). Nests are most often placed in the
top of a large tree (e.g. birch, oak, ginko, ash, pine, elm)

at a height of 4-17m; 50 nests in the Soviet Far East
averaged 8 m above the ground (Winter 1982). There are a
few known instances of Oriental White Storks building their
nests on man-made structures, such as buildings, telephone-
poles and artificial nest-platforms (Sakamoto 1966).
The nest-platform is constructed of dead branches 20-
150cm long and lined with down, dried grass, sedge, and
moss; it is 1.0-2.5 m in diameter and about 0.5-2.0 m high
(Austin 1948:45-46, Austin and Kuroda 1953:333, Winter
1982). Storks often use the same nest-site year after year, if
not disturbed. In the Soviet Far East the foundations of
stork nests are often used as nesting sites by smaller birds,
such as Tree Sparrows (Passer montanus) and White-cheeked
Starlings (Sturnus cineraceus) (Winter 1982). Adults continue
to bring material for the nest even after the young have
become large (Winter 1982, Fei et al. 1991a).
Courtship behaviour has not been fully described in wild
birds. Most observations have been made on captives (see
C.E. King 1988, Archibald and Schmitt 1991). Most of the
described displays seem to be similar to those of the closely
related and well-studied White Stork. Certain aspects of the
Up-Down and Threat Up-Down in the Oriental White
Stork differ in having more side-to-side head and neck
movements, and the head is thrown further back. So far, no
wing-pumping movements have been observed in the
Oriental White Stork however, they might occur in wild
birds when other storks are flying overhead.
It has been reported that the Oriental White Stork's
clattering is of shorter duration than the White Stork's
(Shulpin 1936 cited in Dementiev and Gladkov 1951:455,
Grote 1954, Hornberger 1967:78), conversely it has also
been reported that it is longer (K.M. Schneider 1952, Kahl
1972c, C.E. King 1988:57). Some captive Oriental White
Storks apparently clatter more rapidly than White Storks
(C.E. King 1988:60), whereas others clatter more slowly
(Kahl 1972c). [In both the above cases C.E. King's (1988)
observations were by far the greatest in number.] The
duration and intensity of displays may vary widely, accord-
ing to context and environmental stimuli. Obviously more
study is needed, especially of birds in the wild.
In captives, mutual allopreening, synchronized Up-
Down displays (C.E. King 1990) and cooperative nest-
building (K. Archibald pers. comm. 1990) are indicative of
pair-bonding.
Wild birds copulate on the nest both before and during
egg-laying and after the young have fledged. In wild birds in
China, it has been reported that the female initiates copula-
tion by putting her head under the male's wing (Fei et al.
199la). In captive birds, mates sometimes circled each
other on the nest prior to copulation (C.E. King 1988:96).
The clutch-size is 2-5 eggs. Eggs are milky white to dirty
white and oval to elliptical. In the Soviet Far East, 20 full
clutches in 1975 averaged 3.6 eggs, and in 1976, following
an exceptionally dry winter, 11 clutches averaged 2.73 eggs
(Winter 1982). In captivity, eggs are usually laid at 2-day
intervals (C.E. King pers. comm. 1990). Fei et al. (1991a)
have reported that eggs at a nest in China were laid at
intervals of 4-5 days; however, they gave no details of how
these intervals were determined. There is sometimes as
Oriental White Stork 107
much as a 5 to 7-day difference in age between the chicks in
one nest (Dymin and Pankin 1975).
At a Chinese nest, the birds began incubation 2 days after
the third egg (of 4) was laid (Fei et al. 1991a). Both sexes
share incubation duties and change places on the nest 4-6
times per day (Winter 1982). In one pair (captive), the
incubation duties were shared about equally by male and
female (C.E. King pers. comm. 1990), but in another pair
(wild) the female incubated more and also at night (Fei et
al. 199la). Incubation by wild birds, lasts about 30-34 days
(Austin and Kuroda 1953:333, Fei et al. 1991a). In the
Amur region (USSR), egg shells were observed to be thrown
out by the parents after hatching, and were found on the
ground under the nest (S.W. Winter pers. comm. 1990).
Parental feedings occur about 4-8 times per day, and the
male does most of the feeding when the young are less than a
month old. The bulk of the food given to young in Soviet
nests is fish 80-140 mm long (Winter 1982). Parents re-
ingest any uneaten food (Fei et al. 199la). Young remain in
the nest for 53-60 (Austin and Kuroda, 1953:333), 55-63
(Fei et al. 1991a) or 65-70 days (Dymin and Pankin 1975,
Winter 1982).
After fledging, Soviet young remain within 100-500 m of
their nests for about a week. Then they gather into flocks of
8-40 birds at wetland areas in the general vicinity, before
leaving on their migration in late August (Winter 1982).
Nestlings perform the Up-Down display from 1 day old
onward; the loudness of the clattering increases with age, as
the nestling's bill grows and hardens. Adults were heard to
bill-clatter in response to humans near the nest, and one
female with young performed a Nest-Covering display and
clattered briefly upon sighting the observer near the nest-
tree (Winter 1982).
At Soviet nests, Winter (1982) reported that at 12 nests in
1975, 73.3% of the eggs hatched and 81.8% of the young
fledged. At 11 nests in 1976, 76.7% of the eggs hatched and
91.3% of those young survived for at least 1 week (when
observations ceased). Causes of egg/nestling mortality were
infertility, eggs or small nestlings kicked out of nest by a
parent, large nestlings falling from nest and destruction of
nest by a storm. From 38 nests near the Ussuri River during
1974 and 1975, 62 young fledged, for an average of 1.6
young per nest (Shibaev et al. 1976). Nests are reportedly
sometimes destroyed by Asiatic black bears (Selenarctos thibe-
tanus) (Przhevalsky 1870:196-197, Dementiev and Gladkov
19^1:454, Grote 1954).
TAXONOMY
Whether the Oriental White Stork is a race of its western
cousin, Ciconia ciconia, or whether it is a valid species has
long been debated (Vaurie 1965:85, Kahl 1972c, 1972e,
W.B. King 1981, Neufeldt and Wunderlich 1982, Creutz
1988:18). The two populations are widely allopatric; their
breeding ranges are separated by 50° of longitude, c.
4000 km, and so no tests of reproductive isolation are poss-
ible in the wild. Morphological and behavioural differences
between the two are considerable (C.E. King 1988, Archi-
bald and Schmitt 1991).
108 Oriental White Stork
Because the differences are so great, we have decided here
to treat the two as separate species, although they are
certainly so closely related that they should be regarded as a
superspecies (Amadon 1966).
Morphologically, C. boyciana is much larger than C. ciconia
in body size, culmen and tarsus length. The bill in C.
boyciana is black in adults and greenish with yellow-orange
hues in nestlings, whereas the bill in C. ciconia is red in
adults and black in nestlings. The skin surrounding the eye
and lores in adult C. boyciana is red, whereas in adult C.
ciconia it is black. The iris in adult C. boyciana is pale yellow-
grey to pale blue, whereas in adult C. ciconia it is brown.
There is considerably more silvery grey or white in the
secondaries of C. boyciana.
Behaviourally, in natural breeding areas C. boyciana is
more solitary (i.e. nests are usually 200-300 m or more
apart) and is usually less tolerant of close association with
man than is C. ciconia. Also, in captivity, C. boyciana is much
more nervous and temperamental, needs more solitude and
shows more intraspecific aggression than does C. ciconia.
Noticeable differences have been described in the Up-Down
and Threat Up-Down displays (C.E. King 1988, Archibald
andSchmitt 1991).
CONSERVATION
In the Middle Ages this stork was common in Japan until
1879 and even nested within the city of Tokyo, on top of
Buddhist temples (Austin and Kuroda 1953:332, Yama-
shina 1962b). They decreased drastically during the Meiji
Era (1868-1912), mainly because of illegal hunting. Many
nesting trees of the small remaining population were cut
down during the Second World War. And the few remain-
ing birds were finished off by agricultural chemicals during
the 1950s and 1960s. The last successful breeding in Japan
took place in 1959, when 3 young fledged from 2 nests
(Ornithological Society of Japan 1975:35). Several breeding
attempts during the 1960s produced no young (Fisher et al.
1969:183), and the species has now disappeared entirely as a
breeding bird in Japan.
Although considered a 'good luck omen' by Korean
people (Austin 1948:45), the species has now disappeared as
a breeding bird from that country (Gore and Won
1971:119).
For the Soviet Far East, Winter (1982) lists the following
factors as important in the decline of the species: forest
burning and cutting of large trees used for nesting; drainage
of wetlands and disturbance of hydrological conditions by
construction of power stations upstream; poisoning of food
supplies by chemical pesticides; poaching; and weather dis-
turbances, such as droughts and strong winds. S.M. Smir-
enskii reports (pers. comm. to G.E. King 1989) that recent
droughts and shortage of nesting trees has resulted in low
productivity in storks in the Amur region.
In China, the species is protected by law (M. Williams
pers. comm. 1990). However, hunting and trapping (for
food and for zoos) as well as the conversion of wetlands to
agriculture are threatening the birds (Luthin 1987). Appar-
ently many Oriental White Storks, especially nestlings, are
captured for sale to zoos. Some Chinese zoos sell them to
foreign zoos. At one nest in Heilongjiang Province, China,
only 4 nestlings 'escaped into the wild' during an 18-year
period, most of the others having been poached by humans
(M. Coulter pers. comm. 1990). Nests themselves, as well as
the trees they are in, are sometimes taken for fuel (Fei et al.
199la). Storks wintering in south-central China are killed
by poisons used in hunting wild ducks; they are also threa-
tened by overfishing—which reduces their prey—industrial/
agricultural pollution of feeding areas, and other activities of
man (Wang 1991). Their long bones are sometimes used as
chop-sticks (Dementiev and Gladkov 1951: 454).
This species is currently listed in Appendix 1 of the
CITES Convention, which prohibits international trade. It
is totally protected by law in the USSR, Japan and South
Korea (W.B. King 1981); however, that has not prevented
its extinction in two of these three countries!
Eventually, captive breeding may be important in main-
taining this species, especially if natural populations con-
tinue to decrease. Over 100 Oriental White Storks are
currently (1990) maintained in zoo collections in several
countries. Although this species does not adapt as well to
captivity as does the White Stork, there has already been
some success in captive breeding (C.E. King pers. comm.
1990). Reintroduction programs are highly desirable in
Japan and Korea, if and when it is determined that ecologi-
cal conditions there are suitable for the survival of the birds.
While agreeing that captive breeding is desirable, we
strongly recommend that the taking of young from natural
nests to supply zoos be strictly limited to a few institutions
with facilities and interest in breeding programmes.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 298.
 Blacknecked
Stork
Ephippiorhynchus asiaticus (Latham)

Mycteria asiatica Latham, 1790, Index Ornith., 2, p. 670: India

Other names:
Jabiru, Policemanbird, Green-necked Stork [Australia] (English); Asien-Grossstorch (German);
Banaras, Loharjang, Loha sarang (Hindi); Lohar jangh, Ram salik (Bengali); Telia-hereng
(Assamese); Periya-narai (Tamil); Pedda nalla konga (Telugu); Ali-kokka (Sinhalese): Hnet-kalah
(Burmese)

IDENTIFICATION
The Blacknecked Stork, one of the tallest, most stately and sedate of the storks, stands 129-150 cm tall.
A slim black and white bird with a long, thin, slightly upturned black bill and very long bright coral-red
legs. Its large size is well described by its Sinhalese name which means 'man-heron'.
The head, neck, scapulars, greater and median wing-coverts, and tail are glossy black, showing
purple, green and blue iridescence according to the angle of the light. The back, flight feathers, lesser
wing-coverts and underparts are pure white. When spread, the white wings have a diagnostic black
band, running longitudinally down their centre, both top and bottom. The large bill is a shiny black.
The sexes are easily separated by eye colour: the male's is brown, the female's bright yellow and
contrasts sharply with the dark feathers of the head; furthermore, males are noticeably taller. [Some
published accounts, such as Andersson's (1872:282), have the eye-colour of the sexes reversed.]
The most noticeable change from the regular adult plumage to the breeding plumage is that the legs
become brighter. The gloss on the black portion of the plumage is heightened during the courtship
period. And there is a 'skull-cap' glossed crimson-purple or bronze-purple (Iredale 1956:112, Henry
1971:382).
The small, downy young are snow-white at first, the head and neck changing to dusky grey later.
Their bills are straight, not upturned as in adults (Gates 1883:264, Lucas and Le Souef 1911:127,
Lowther 1949:pl. 58, S. Ali and S.D. Ripley 1968:104). Immature birds are dull brown with whitish
underparts, lower back, and rump; they have flight feathers tipped blackish brown and basally white
and dull blackish legs. The white parts including the belly are dusky brown, turning white with age.
Juveniles are patterned like adults, but the glossy black areas of adults have a brownish cast in young
birds. We have seen fully adult plumed birds with legs still dark brown; such birds are probably
subadults. Immatures of both sexes have brown eyes (Dharmakumarsinhji 1955:93).
Vocalizations include bill-clattering loudly during Up-Down and other displays (Kahl 1973b). The
species is said to make 'low grunts' at the nest (Beehler et al. 1986:60) and to utter 'dull booms' (Pizzey
1980:62), but we find these statements doubtful.
'In flight it is a noble sight with its neck extended, its long, bright red legs projecting behind the tail,
and the huge white wings marked with a broad black band formed by the greater and median coverts;
this pattern is repeated on the underwing' (Henry 1971:382). The flap rate was measured at 167/min
(n = 20) by Kahl (1971b). The Blacknecked Stork looks 'rather skeletal' in flight (Pizzey 1980:62).

This species is unlikely to be confused with any other bird
in its range. During the winter months, in India, it could be
mistaken for a Black Stork, but the latter species is smaller,
all dark above, and has a red bill. In Australia, there is a
slim possibility that it might be confused, at a great dis-
tance, with a Magpie Goose (Anseranas semtpalmata),
although the latter is much smaller and flies with a faster
wing-beat.
DISTRIBUTION AND POPULATION
This species formerly ranged throughout the entire Indian
subcontinent, from Sri Lanka southeastward through
Burma, Thailand to the northern Malay Peninsula, Kam-
puchea, southern Laos and Vietnam. It also occurs disconti-
nuously in southern New Guinea (Irian Jaya and Papua);
islands in the Torres Strait; and northern, northwestern and
eastern Australia. Occasionally it wanders south as far as
southern Victoria, Australia (Bell 1963, Beruldsen
1980;155, Blakers et al. 1984:62). Accidental (once: 1898) in
western Java (Kuroda 1936:541).
The birds appear to be sedentary, and we know of no
evidence of regular long-distance migration. In Australia, it
wanders far from its breeding grounds but there is nothing
to suggest regular seasonal movements (Blakers et al.
1984:62).
Solitary nesting habits limited the species' numbers in
suitable habitat, and historically it was nowhere abundant.
However, within the last two decades, numbers on the
Indian subcontinent have fallen dramatically. In Pakistan it
is very rare and no longer resident, being only an occasional
straggler to this country (TJ. Roberts pers. comm. 1988).
In India, it remains widespread but thinly spread: the states
of Rajasthan, Uttar Pradesh and Assam seem to be the
present stronghold, and the greatest declines have come in
Blacknecked Stork 111
Bihar and West Bengal; it was never common in the south
(Rahmani 1989a). In Gujarat we have seen non-nesting
pairs of birds in areas where traditional nest trees have been
destroyed, and we suspect that failure to breed may be
because of lack of alternative suitable sites (pers. obs.
J.A.H.). In the Keoladeo National Park, Rajasthan, Kahl
(1970:457) found 5 nesting pairs in 1967, but we have
recently seen only 2 pairs and these are dependent on water
supplies that are subject to poor monsoon periods. During a
mid-winter census (Dec 88-Feb 89), 120 were counted in
India and 4 in Sri Lanka (D.A. Scott and P.M. Rose
1989:56) [It is apparently an error that 63% of those were
listed for the state of Tamil Nadu (D.A. Scott and P.M.
Rose 1989:43), for Rahmani (1989a:104) states that there
has been only one record for the entire state in recent years.]
In Bangladesh, the species has disappeared (M.A.R. Khan
1987:230), though it is still an occasional visitor to Nepal
where it sometimes nests in small numbers. In Sri Lanka, a
relict population of 6-7 pairs persists in the Yala National
Park (Rahmani 1989a:105).
Elsewhere, scattered birds have been reported from
Burma and Thailand, but there are no reports of recent
nesting (P. Round pers. comm. 1989). Four birds were still
present in the Hat Chao Mai National Park, near Trang,
southern Thailand, in December 1980 (P. Round et al.
unpub. report), where it was once found breeding (on cliffs)
in 1917 (Medway and Wells 1976:96). It is believed that a
few pairs remain in the Mekong Delta in Vietnam in the
Melaleuca forests (Le Dien Due pers. comm. 1987) and
possibly in Cambodia (V. Quy pers. comm.). It is thought
that the species has been in decline for several decades due
to increased cultivation, especially for rice (Luthin, pers.
comm. 1988), and we confirm from our own observations
over four decades that this is so.
Only in southeastern Irian Jaya (Indonesian New
Guinea), where an estimated 650 birds were counted during
a dry-season concentration (Bishop in Silvius and Verheugt
1990), and in Australia does this species seem to still
prosper. In Australia, it is widespread and common in the
north. It breeds in the Kimberleys, Cape York and the
Atherton regions and in the swampland areas of the Top
End. We have seen it in some numbers on the flood plains of
the South Alligator River. Further south it occurs in
swamps associated with large river systems. In winter, there
is some movement to the Pilbara and Murray-Darling re-
gions. The only population estimates from Australia are 37-
43 birds in New South Wales in 1965 (Blakers et al.
1984:62).
ECOLOGY
The species appears to require freshwater swamps, rivers,
lakes with shallow water, coastal areas and river estuaries
together with suitable large trees, or at least one large tree,
some distance away. Most pairs used the same territory
continuously, but where such areas dry out intermittently,
birds may wander to other areas that continue to provide
food. A lone escaped bird that survived in the Everglades
(Florida, USA), for 4 years, followed a similar pattern,
112 Blacknecked Stork
remaining in one local area for most of the year but retreat-
ing to deeper water some 30 km away in the dry season
(pers. obs. J.A.K.).
This tall species typically forages in shallow water, Walk-
ing Slowly, with measured strides, at about 1 m/sec, and
Probing in the water and submerged vegetation. The bill is
held open about 75-100 mm at the tip and raised about
200 mm above the water between probes (Kahl 1973b, pers.
obs.).
Food location seems to be mainly tactile but sometimes
also visual. Occasionally individual birds Run a short dis-
tance in pursuit of active prey, and we have seen this quite
frequently in Australia. The escaped bird observed in the
Florida Everglades fed by Standing and Walking, and in-
frequently captured fish (pers. obs. J.A.K.). A bird watched
feeding for 2 h at Kakadu National Park, northern Austra-
lia, stalked forward at about 1 step/s with the bill held open
and jabbing vertically into the water as deep as the junction
of the mandibles. The bird caught many small fish, bit them
several times in the tip of the bill, and then swallowed by
raising the bill to the horizontal and tossing the head back.
Food location seemed to be non-visual, as the bird jabbed in
the water at regular intervals while wading, without appear-
ing to spot any prey. It crossed and re-crossed an area about
20 X 50 m in size within a large marsh. In the 2 h period the
bird was seen to capture only 10-15 small (25-75 mm) fish,
although it made an estimated 2000-3000 probes in the
water. Another bird, also in Kakadu Park, was seen and
heard foraging almost constantly for over 4 h (02.00-
06.15h) on a clear, starry and moonless night. It Probed
deeply and forcefully into the water, making a loud splash
with each plunge, at intervals of just under 1 s, as it waded
silently back and forth in a small billabong just 10 m from a
tent (pers. obs. M.P.K.).
Pairs forage independently but often within sight of each
other; if one bird takes wing, the other will usually follow
within a few seconds. We have seen small parties feed quite
closely together, usually consisting of a pair of adults and
juveniles. Despite this communality in family groups, the
birds are normally highly territorial during the nesting
season. In one instance, two birds rose into the air, chasing
each other, the dominant bird grasping the wing of the other
in midair in a spectacular aggressive display. In the Keola-
deo National Park, India, we have seen a bird land amongst
a party of feeding egrets and drive them from the area (pers.
obs. J.A.H.). At the same place, a pair, accompanied by a
large, but still Begging, young bird, threatened and attacked
a flock of White Pelicans (Pelecanus onocrotalus) at a feeding
area (Breeden and Breeden 1982).
Drinking is remarkably frequent, and Kahl (1970, 1973b)
reports one bird drinking 21 times in succession by placing
the open bill in water and scooping upward and forward to
above the horizontal to swallow.
Kahl (1973b) described a Flap-Dash display, usually
seen at feeding areas. One bird, usually the male, stands
erect then runs with long strides and flapping wings. It
dashes away from its mate, then dashes back and stands
with wings widely spread. This is probably a social display,
making use of the flashing white wing pattern, in pair-
formation and maintenance of the pair-bond. The Flap-
Dash has been reported only from the Blacknecked, Saddle-
bill and Jabiru Storks, also the only species with large areas
of white in the wing.
Prey consists mainly offish, but also frogs, reptiles, crus-
taceans and large insects as well as other small animals
(Rand and Gilliard 1967:48). Breeden and Breeden (1982)
saw a Blacknecked Stork catch, kill and fly off with a Pond
Heron (Ardeola grayii) at Bharatpur, India. They have also
been observed catching and swallowing whole live Coots
(Fulica atra) (Panday in S. AH and S.D. Ripley 1968).
Blacknecked Storks are usually solitary or in pairs, but in
Australia, they have been found in parties of up to 18
(Blakers et al. 1984:62). The flock of 50-60 reported in India
(Whistler 1918a) was probably an error (A. Rahmani pers.
comm. 1990). Earlier this century, it was said that Cambo-
dians prize their blood as a medicine (Delacour 1929, Dela-
courandjabouille 1931:88).
BREEDING
Most nests in India are 6-25 m above ground and have been
found in trees such as kadamb (Acanthocephalus kadamba),
peepul (Ficus religiosd), simul (Bombax malabarica) and shees-
hum (Delbergia sissoo); in Australia, nesting is in trees such
as Eucalyptus spp., Nauclea orientalis and Melaleuca spp. Con-
struction of a new nest may take more than a month (Hume
and Oates 1890:266). It is built by both birds and often
reused annually. This species prefers nesting in a high,
isolated tree, which presumably allows easy access for its
large wing-span. One nest was built on cliffs, on Telibun
Island off the west coast of peninsular Thailand, near Trang
(H.C. Robinson and F.N. Chasen 1936:208, Madoc
1950:57).
In various parts of its range, egg-laying seems to be
mainly as the rains cease (P. Slater 1959). In northern
India, this is usually in October; if the rains end earlier than
normal, the birds may lay in September (Hume and Oates
1890:267).
Pairing is long-term, thought to be for life, so that when
building or rebuilding commences there is less demonstra-
tive behaviour than in stork species that pair anew each
year. The greeting ceremony, which is not often seen, con-
sists of Up-Down displays (Kahl 1970, 1973b) in which the
mates face each other with wings fully open and fluttering
rapidly in a horizontal arc. Loud clattering occurs 8-
10 times/s for 5-10 s. Less commonly the Up-Down may be
performed away from the nest, at the feeding grounds
(Hume and Oates 1890:266, Barrett 1954:64).
The Erect Gape can occur either at or near the nest,
shortly before flight; it is similar to the homologue in the
Marabou Stork, but no sound is uttered. A single snap or a
short series of bill rattles can take place during the Arching
display, which occurs when the birds are disturbed at the
nest. Clattering by the male does occur during copulation,
which starts for the most part without obvious preliminar-
ies, except that sometimes the female presses up against the
standing male. During Copulation Clattering, the female's
wings are partially or wholly open, and the male flaps his

wings slowly for balance. Copulations average 24 s in dur-
ation (range = 15-32, n = 4).
Both sexes share in building the nest. The nest is a
massive structure—usually 1-2 m in diameter and 1.0-
1.5 m deep, with a considerable cavity—built high in a large
tree, made of sticks and lined with straw, leaves, rushes,
grass, sometimes mud and often rags.
The clutch size is 3-4, rarely 5. The eggs are broad ovals,
compressed at one end, and dull white with the inner mem-
brane dull green. Eggs become soiled and stained after a few
days in the nest and become a dingy yellowish white (Hume
and Gates 1890:268, S. Ali and S.D. Ripley 1968). Parents
at nests in India were seen regurgitating water over the
contents of nests presumed to contain eggs (Kahl 1970).
Both sexes share in incubation, but the incubation period
is unknown. Both parents feed the young by regurgitation
onto the nest floor (S. Ali and S.D. Ripley 1968). Frith
(1976:86) mentions that parents give water to the young on
hot days. However, he does not give a reference for this
information, and it may have come from a mis-reading of
Kahl (1973b), which is not very clear on this point. Kahl
(1973b) did not observe the young of the Blacknecked Stork
but, instead, recorded Saddlebill parents watering young.
The fledgling period is given as 100-115 days (Frith
1976:86). After they can fly, young still solicit food from
their parents away from the nest (Breeden and Breeden
1982).
TAXONOMY
The Blacknecked Stork was formerly placed in the mono-
typic genus Xenorhynchus\ however, it is so obviously closely
related to the Saddlebill Stork (Ephippiorhynchus senegalensis)
that it makes good sense to combine the two into one genus
(Kahl 1970, 1972e). Plumage differences from E. senegalensis
are minor. In E. asiaticus the lesser and median under wing-
coverts are white and the uppertail-coverts are white, whereas
in E. senegalensis most of the lesser, all of the median and the
greater underwing-coverts are black (only a narrow band of
underwing-coverts on the leading edge of the forearm are
white) and the uppertail-coverts are black (Sharpe 1898:311,
313; pers. obs. M.P.K.). Other morphological differences
involve mainly the coloration of the bill and legs.
Two subspecies have been described and were previously
Blacknecked Stork 113
recognized—E. asiaticus asiaticus (Latham) and E. australis
(Shaw) —but having studied them both in the field and in
museum trays, we cannot separate the populations. Suppos-
edly the New Guinea/Australian birds have a greener gloss
on the neck (Iredale 1956:112). However, birds we have
observed in both India and Australia appeared the same,
and the amount of blue or green iridescence on the neck
varied with the quality of the ambient light. Amadon and
Woolfenden (1952) and Legge (1880:1116) also found the
Australian birds difficult to separate from those on the
Asian mainland. Therefore, we do not recognize any sub-
specific division.
CONSERVATION
Habitat destruction, in the form of cutting of traditional
nesting trees and draining of wetland feeding areas, is an
inevitable consequence of the growing human populations
throughout Asia. Blacknecked Storks are solitary and highly
territorial—at least during the breeding season—and suc-
cessful reproduction means little to the overall population if
offspring have no place to go.
Trapping for zoos, both foreign and Indian, is given as
another likely cause of some of the drastic decline in India in
recent years (Rahmani 1989a). Overfishing competes for
food supplies, and pesticide poisoning is a possible threat in
highly agricultural areas. Shooting is probably not a serious
concern, because the flesh of the Blacknecked Stork is too
fish-tasting to eat (A. Rahmani pers. comm. 1990).
For the conservation of this species in the Indian subcon-
tinent, Rahmani (1989a) lists the following six steps: (1) a
survey to identify important wetland breeding sites; (2)
protection of critical habitat; (3) legal protection of the birds
themselves; (4) ecological research; (5) captive breeding;
and (6) publicity, to gain public support.
The Government of India has recently placed the Black-
necked Stork in Schedule 1 of the Indian Wildlife (Protec-
tion) Act, and its trapping has been banned.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 299.
 Saddlebill Stork
Ephippiorhynchus senegalensis (Shaw)

Mycteria senegalenis Shaw, 1800, Trans Linn. Soc. London, 5, p. 35, pi. 3: no locality (=Senegal)

Other names:

African Jabiru (English); Jabiru de Senegal (French); Sattelstorch (German); Saalbekooievaar

(Afrikaans); Abu Mijeh, Abu Milieh (Arabic); Hello (Mandingo); Baba da Jika, Zuguntumi
(Hausa); Kandjendje (Kwangali); Hukumihlanga, Kokwasabi (Tsonga); Kololoane (Chuana);
Nyamantando (Sotho)

IDENTIFICATION
The Saddlebill Stork is one of the tallest of the storks, standing at 135-150 cm. A slim, sedate, and
stately black and white bird, it has a long, thin, slightly up-turned, multi-coloured bill and very long
legs.
The adult's head and neck are black, and the underparts white. The white wings have a diagnostic
black longitudinal band, above and below. The black head, neck, wing scapulars and tail- are glossy,
showing purple or green according to the angle of the light. The tail- and uppertail-coverts are black.
The large, slightly recurved bill is alternately red, black, red; the distal half (c. 15 cm) is deep red, the
middle third (c. 10cm) is black, and the basal fifth (c. 6cm) is covered by a deep red fleshy lappet,
which extends back to the eye. Above this red lappet is a bright yellow lappet, forming the 'saddle'. The
legs are blackish to greyish green, with pink tibio-tarsal joints and toes. Males are taller and have a dark
brown iris, whereas females have a pale yellow iris [erroneously reversed in Andersson (1872:282)].
Breeding plumage is not known to differ noticeably from regular adult plumage, except that the
iridescence on the black regions of the head and neck is more pronounced. One or both members of the
pair often have two small yellow, sometimes red, wattles that hang down 15-35 mm under the base of
the bill, like 'stirrups' under the 'saddle' (Kahl 1973b; Brown et al. 1982:185). A small (45-50 mm
diameter), round or oval-shaped patch of bare, dark red skin is sometimes exposed in the middle of the
breast (Chapin 1932a:462); it is often hidden by feathers. These colourful areas of bare skin may
indicate fully mature birds in breeding condition.
Details of the development of nestlings are minimal. The downy young are fluffy and white, the head
and neck later changing to a darker, smoky grey. The bill is horn-coloured and 'not particularly long'
(Mackworth-Praed and Grant 1962); by 21 days the bill is still only 88mm long (Pitman 1965). At
45 days, nestlings are still largely downy, with dark feathers showing on the wings (Brown et al.
1982:186). The plumage of large nestlings is mainly a 'lustreless greyish black; the head and neck are
covered with grey down; the bill is blackish, with only a slight indication of red at the tip; the "saddle"
at the base of [the bill] is blackish and edged with a fringe of whitish down' (A.L. Butler 1905). Other,
captive reared, young still did not show red on the bill at 6 months of age (F. Junor, pers. comm. 1965).
Soon after fledging, young still have the head and neck covered with thick and woolly, snuff-coloured
down (Niven and Niven 1966b).
Juveniles show little or no red in the bill or legs; areas that are black in adults are dark grey; areas
that are white in adults are pale grey (C.W. Benson and C.R.S. Pitman 1958). By the second year, the
bill is dull red and black, and the 'saddle' is a dull yellow (Brown et al. 1982:185).
Vocalizations, if any, are unknown in adults, although they do bill-clatter during some displays, as
does the closely related Blacknecked Stork of Asia. Begging calls of nestlings are less loud than those of
the colonial storks (Kahl 1973b).
116 Saddlebill Stork
An impressive bird on the wing, with large areas of white
in the flight feathers and a span of over 2.5 m, it often soars
on thermals until nearly out of sight from the ground. The
flap rate was measured as 150/min (n = 13) (Kahl 1971b).
It flies with the neck and feet fully extended, and, because of
its extreme length, 'it flies in a curiously bent position with
head and feet lower than the body' (Mackworth-Praed and
Grant 1962:83).
The tallest African stork, the Saddlebill is slimmer and
more graceful than the Marabou. Juveniles, in grey plu-
mage, might be mistaken for juvenile Yellowbilled Storks
but are much taller. Saddlebill juveniles, if not in the com-
pany of their parents, might possibly be misidentified as
young Marabous (Turner in Pitman 1965).
DISTRIBUTION AND POPULATION
This species occurs throughout tropical Africa, from Sene-
gal to Sudan and Ethiopia, south to northern Namibia,
northern Botswana and northern South Africa (Transvaal,
Zululand). It occurs sporadically further south but is quite
rare south of the Orange River.
The birds appear to be sedentary, and we know of no
evidence of regular long-distance migration. They are to
some extent nomadic, especially during drought conditions
or when the large rivers are in flood (Chapin 1932a:463,
Bannerman 1953:177).
The Saddlebill Stork is widespread but never very numer-
ous. Its territoriality and solitary nesting limits numbers in
suitable habitat, and historically it was nowhere abdundant.
There are 6 recorded breeding sites in Kenya, 5 in Uganda,
and 11 in Tanzania (Kasoma and Pomeroy 1987). The
record of 100 birds at Lake Turkana, Kenya (Kasoma and
Pomeroy 1987) is apparently an error. Along the 52km
length of the Kazinga Channel, in western Uganda, only 1
pair of Saddlebills was found over an 8-month survey
(Kasoma and Pomeroy 1987). In the Kafue flats, Zambia,
Dowsett and de Vos (1965) counted an average of 27 Sad-
dlebills in an area of 58 square miles (=151 km 2 ), 1 bird/
5.6 km2; further northeast in the Bangweulu Swamps, Zam-
bia, G.W. Howard and D.R. Aspinwall (1984) estimated a
density of 1 bird/5.8 km2. In South Africa, where the species
occurs regularly only in Zululand and the eastern Trans-
vaal, Siegfried et al. (1976) estimate a total population of
'fewer than 50 birds*.
ECOLOGY
The Saddlebill Stork is found in aquatic habitats in open or
semi-arid country, generally avoiding deep forest. It favours
shallow freshwater marshes, rivers, lake shores and flood-
plains, with suitable large trees, often some distance away,
for nesting. Most pairs use the same territory continuously,
except during drought, when the birds may wander nomadi-
cally.
Birds of this tall species typically forage in shallow water,
Walking Slowly and Probing in the water and submerged
vegetation. They usually take 0.7-1.0 steps/s and each
stride covers 0.8-L2 m. The bill is held open about 50-
70 mm at the tip and raised about 200 mm above the water
between probes (Kahl 1973b). There is some evidence that
large fish may be impaled with one of the mandibles, rather
than just grasped (Bell-Cross 1974). Most prey finding
appears to be tactile. However, occasionally individual
birds Run a short distance, often with wings open for bal-
ance, in pursuit of active prey. Foot Stirring has been
reported (Brown et al. 1982:185). Birds have been seen
washing frogs in water before eating them (Curry-Lindahl
1961).
Pairs forage independently but usually within sight of
each other. Small parties sometimes feed quite close
together and usually consist of an adult pair and their
offspring. Despite the communality in family groups,
Saddlebills are usually highly territorial (Brown et al.
1982:186); however, this territoriality may become weaker
outside the breeding season (G.W. Howard and D.R.
Aspinwall 1984). Most foraging is diurnal, although a bird
has been reported feeding at night at an artificially lighted
waterhole at Mulika Lodge, Meru National Park, Kenya
(Lassus 1973).
Kahl (1973b) described a Flap-Dash display, usually
given at feeding areas. The male (usually) stands erect, runs
away from its mate with long strides and flapping wings,
turns and runs back to the mate, and stands with wings
widely spread. This is almost certainly a social display-
making use of the flashing white wing pattern —and prob-
ably functions in maintenance of the pair-bond, or even in
pair-formation.
Prey consists mainly offish, but also frogs, reptiles, crus-
taceans, some molluscs and other small animals. Where
they are common, lungfish (Protopterus aethiopicus) are favor-
ite food (F J.Jackson 1938:76, Kahl 1973b). Adults com-
monly swallow fish 15-30 cm in length and weighing up to
500 g, occasionally 1300 g. They have also been reported
eating large water beetles (Chapin 1932a:463), other insects

and carrion (Heuglin 1869-73:1113). Prey is often washed
before swallowing head-first; and birds commonly drink
after eating (Brown et al. 1982:186). Drinking is frequent,
with the bird placing the open bill pointed forward in the
water and scooping upward and forward to above the hori-
zontal to swallow. Snipping off catfish spines before swal-
lowing the fish has been seen (A. Morris 1979). Scavenging
on fish regurgitated by cormorants has been reported
(Kasoma and Pomeroy 1987).
This species usually occurs solitarily or in pairs. Small
groups seen are often parents and several young (C.W.
Benson and C.R.S. Pitman 1958). The largest recent aggre-
gation that we know of is a group of 12 birds seen in Zambia
(G.W. Howard and D.R. Aspinwall 1984), although Ban-
nerman (1953:177) reported 'flocks of twenty to thirty may
be observed at times on the sandbanks' and 'one record of
thirty seen on the Upper Niger'. Curry-Lindahl (1961)
reported a pair of Saddlebills that stayed near a herd of
hippopotamuses and moved to the centre of the herd when
humans approached.
Saddlebill Storks roost on trees. Normally they are rather
shy but they can become tame in areas such as national
parks where they become accustomed to vehicles (Brown et
al. 1982:185).
BREEDING
At the northern and southern ends of the range, Saddlebills
normally begin nesting late in the wet season or at the start
of the dry season; in equatorial regions they may nest during
the rains. In both cases, the young usually fledge at the
height of the dry season, when food is concentrated and
relatively easy to obtain.
The nest is usually built in the top of a large tree (e.g.
Euphorbia ingens, Euphorbia dawei, Acacia giraffae, Kigelia pin-
nata, Adansonia digitata). Especially in Zambia, they often
build nests in trees (e.g. Albizzia harveyi) growing out of
termite-mounds (C.W. Benson and G.R.S. Pitman 1958).
Unless in a well-protected area such as in an inaccessible
swamp, the nest is often as high as 20-30 m from the ground
(Brown et al. 1982:186). They have been reported as some-
times nesting on cliffs (Mackworth-Praed and Grant
1962:83, Clancey 1964a:46), but this needs confirmation
(Pitman 1965).
Saddlebill Storks are reported to occasionally use old
nests of other large birds, such as Tawny Eagles (Aquila
rapax). Sometimes other large birds, such as Secretary Birds
(Saggitarius serpentarius), usurp unoccupied Saddlebill nests
as well (PFIAO nest-record cards, University of Gape
Town). A SaddlebilPs nest in western Kenya was sur-
rounded, on two consecutive seasons, by many nests of
Blackheaded Weavers (Ploceus cucullatus) that arrived after
the storks began to nest (Kahl 1973b). These smaller birds
probably gain considerable protection from predators by
establishing their colony near a Saddlebill's nest (Moreau
1942).
Pairing is long-term, thought possibly to be for life, so
that when nesting starts the birds are less demonstrative
than stork species that mate with different partners each
Saddlebill Stork 117
year. Sometimes many days pass during the breeding season
without any obvious social displays taking place.
This is the only species of stork for which the Up-Down
display has not been adequately described in nature. It was
not seen at a Kenyan nest in 146 h of observation over two
seasons (Kahl 1973b). Professor H. Mendelssohn (pers.
comm. 1972) observed the Up-Down in a pair of captive
Saddlebills at the Tel-Aviv Zoo. In this pair, the period of
display was prolonged, because the birds broke their eggs
and had to re-lay. They gave Up-Downs, as a greeting
when one bird was on the nest and its mate approached:
The approaching partner came with quick and very
long steps, head and neck stretched forward. Arriving
at the nest, he (or she) spread his wings fully and began
a quick bill-clattering, lowering the head and some-
times lifting it again to [the] horizontal position and
even higher. Then the wings were closed, the head
lowered and the bird generally pecked at sticks in the
nest. The partner on the nest displayed similarly, but
less intensively, beginning after its mate. The wings
were less widely spread, sometimes not at all.
Other social displays are detailed and illustrated in Kahl
(1973b). In the Arching display, which often occurs when
the birds are disturbed, the stork stands erect, leans forward
and compresses its plumage; a loud snap or a short series of
bill-rattles may be given. The Erect Gape occurs on or near
the nest, usually before flight; it is similar to a homologous
display in the Leptoptilos storks, but no sound is uttered by
the Saddlebill. Copulation has not been described in the
Saddlebill.
We once observed a Forward Display by a captive imma-
ture Saddlebill as it threatened several Yellowbilled Storks
in its feeding-pond. The Saddlebill stood erect, spread its
wings, and clattered its bill slowly, as it approached the
smaller storks (Kahl 1973b). A similar display was given, as
a threat, to humans by a captive adult in the Tel-Aviv Zoo
(H. Mendelssohn pers. comm. 1972). The bird approached
with widely spread wings and lunged with quick bill-
clattering. Frank Junor (pers. comm. 1965) described how
captive-reared juveniles, once they could fly, 'would often
fly low over the head of a human, lower their heads in flight
and snap with a most audible clack above the head of the
stranger'.
This species does not necessarily breed every year in a
given locality (Kemp 1974:15, Brown et al. 1982:186). It is
sometimes said to be shy and nervous at the nest (Pitman
1965), but a pair near Kisumu, Kenya, nested in two suc-
cessive years in a tree within 20 m of a busy, tarmac high-
way (Kahl 1973a).
Both adults cooperate in building the nest, but at a Kenya
nest the male brought most of the sticks. The platform is a
massive structure made of sticks and lined with reeds,
sedges, or mud. It is built or repaired by both birds and
commonly reused year after year. Nests may be up to 2 m in
diameter and 50 cm deep (occasionally larger), with a cen-
tral depression deep enough to completely hide a sitting bird
(Pitman 1965, Brown et al. 1982:186).
The eggs are oval, dull white, slightly glossed and coarse-
118 Saddlebill Stork
textured (Pitman 1965). The clutch size is 1-5, usually 2-3;
the mean of 54 clutches was 2.8 eggs (Brown et al.
1982:186). According to Pitman (1965), the clutch size
north of the Equator is usually 3 eggs, whereas in Zambia
and Zimbabwe clutches of 4 are more frequent.
Both sexes incubate, in shifts that vary from 1 h 25 min to
5 h 50 min. The incubation period has not been accurately
recorded, but has been estimated at 30-35 days (Brown et
al. 1982:186). Old birds often sit in the nest on their tarsi
(Heuglin 1869-73:1113). Parents often bring water back to
the nest and regurgitate up to 1-2 litres onto the nest and
eggs(Kahl!973b).
Both parents feed the nestlings by regurgitation onto the
floor of the nest. Young are also given water by their
parents, who stand upright, hold their bills high, and regur-
gitate it over them (Kahl 1973b). Young perform Begging
displays, either standing or on their tarsi, with rhythmic
'up-down' nods of the head, slightly opened wings and
rather weak braying vocalizations; they sometimes bill-
clatter to the parent. Unattended nestlings threaten
intruders with a Nestling Bristle display, which includes
erection of head, neck, and upper back feathers, and clatter-
ing of the bill (Kahl 1973b).
Chicks are brooded closely for the first 10 days. The
fledgling period is not accurately known but is probably at
least 100 days (Kahl 1973b, Brown et al. 1982:186); this is
considerably longer than Pitman's (1965) estimate. Fledged
young still give Begging displays and are fed by the parents
after they leave the nest (Niven and Niven 1966b). At what
age juveniles become fully independent and separate from
their parents is not known; however, it is probably several
months after they leave the nest. Family groups are often
seen together long after the breeding season, and, according
to Pitman (1965), 'they are still together when the red
colour is appearing on the bill [of the juveniles] and when
the yellow "saddle" begins to show'.
Nesting success has not been recorded. 1-2 young nor-
mally fledge from a clutch of 3, apparently more in southern
Africa (Pitman 1965, Brown et al. 1982:186). Sample counts
at Lochinwar and Blue Lagoon, in Zambia, during August-
November at the end of the breeding season showed 'no
more than 5% immature birds' (Dowsett and de Vos 1965).
TAXONOMY
The Saddlebill Stork (genus Ephippiorhynchus) is so closely
similar to the Blacknecked Stork (formerly classified in the
monotypic genus Xenorhynchus) that they have been com-
bined (Dekeyser 1962, Kahl 1972e).
Plumage differences from E. asiaticus are minor. In E.
senegalensis most of the lesser, and all of the median, and all
of the greater underwing-coverts are black (only a narrow
band of underwing-coverts on the leading edge of the fore-
arm are white) and the uppertail-coverts are black, whereas
in E. asiaticus the lesser and median underwing-coverts are
white and the uppertail-coverts are white (Sharpe 1898:311,
313; pers. obs. M.P.K.). Other morphological differences
involve mainly the coloration of the bill and legs.
CONSERVATION
The population is probably stable but could easily become
threatened (Brown et al. 1982:185). The taking of nestlings
for sale to zoos could make an impact on a bird that nests
solitarily and is nowhere abundant, such as the Saddlebill.
In 1989, the Dallas Zoo (USA) paid US$6000-6500 per bird
(P. Osten pers. comm. 1990).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 300.
 Jabiru Stork
Jabiru mycteria (Lichtenstein)

Ciconia mycteria Lichtenstein, 1819, Abhandl. K. Akad. Wissen. Berlin, Phys. Kl. (1816-17), p. 163;
(based on 'Jabiru' of Marcgrave, 1648, Hist. Rerum Nat. Brasiliae, p. 200): Brazil.

Other names:

Garzon soldado, Garzon bianco, Cigiiena de cuello pelado, Galan sin ventura, Juan grande, Baguari
(Spanish); Blaasman (Dutch); Doih (Warrau); Tararamu (Macusi); Mora-coyasehre (Arawak);
Tuyuyu (Guaram); Tuiuiu-corol, Jaburu-moleque, Jaburu (Portuguese)

IDENTIFICATION
The Jabiru Stork is a robust, all white-plumaged stork, standing 120-150 cm tall and having a black
head and black, massive, slightly upturned bill. The common name 'Jabiru' apparently derives from the
South American Indian word 'Yabini', meaning 'blown out with the wind', and referring to the loose,
brilliant red skin of the lower neck, which becomes inflated during threat, anger or courtship.
This is the only stork with all-white feathers, including wings and tail. The head and neck are
featherless, except for a silvery grey tuft of feathers on back of the crown. The upper neck is dull black
skin that abuts a band of loose pink skin, forming an inflatable throat sac around the lower third of the
neck; it inflates and turns fiery scarlet during anger, excitement or physical exertion (P.L. Sclater and
W.H. Hudson 1889:106, Kahl 1971e, 1973b). A small oval-shaped patch of bare skin also occurs on the
chest; although normally covered with feathers, it becomes exposed when the bird stands erect, as
before take-off or during displays. The flight feathers (both primaries and secondaries) are white,
making it the only all-white species of stork. The legs and feet are black, and the iris is brown in both
sexes. The sexes have similar plumage, although the females are slightly smaller and have thinner and
more upturned bills (Kahl 1971e, B.T. Thomas 1985).
Individual birds can be distinguished by variations in the configuration of the black/red junction on
the bare neck (Sick 1985, B.T. Thomas pers. comm. 1990). Some birds even have small spots of red
within the black or black within the red. Such differences, which seem not to change as the bird ages,
may be helpful to researchers studying birds that cannot be individually marked.
Plumage, which is often soiled at other times of the year, moults into immaculate white in the
breeding season. The bare skin of the neck is smooth and silky to the touch (B.T. Thomas 1985). The
red band on the lower neck brightens and, because of increased sexual excitement, is a rich scarlet much
of the time.
Hatchlings are covered—except for the naked lores and orbital skin—with a greyish white down, with
localized yellowish tinges, which is especially thick on the head and uppersides, and more scantily
covered on sides and undersides. They have straight, rather than recurved, bills (Lloyd 1895, Bent
1926:69).
Juvenile plumage is mottled brownish grey, the feathers being light grey with darker tips. A sparse,
dusky brown, hair-like down is retained on the head and neck, being longest on the occiput where it
forms a blackish crest. The undersides are mostly white, and the white wing feathers tipped with grey or
buffy brown. The remainder of white plumage develops gradually over the first 2 years, beginning at the
undersides and above the tail (Nutting 1882, Hagmann 1907, Bent 1926:69).
Generally this species is voiceless, but it bill-clatters loudly in displays and in alarm. A coughing
sound is occasionally given in Up—Down greeting displays and during copulation.

Having the second-longest wingspan of any New World
bird-'nearly eight feet' ( = 2.4m) (Hudson 1920:117)-the
Jabiru flies gracefully, with legs and neck extended, the
latter appearing disproportionate because of a bulge formed
by the loose skin on the neck (Roosevelt 1914:91). This
bulge gives the appearance of flying with the neck partially
retracted. Taking off requires a running start. The bird
alternates a few slow flaps with gliding, and frequently uses
thermals for soaring (Chubb 1916:159, Wetmore 1926:60,
Sick 1985). Dickey and van Rossem (1938:87) commented
that 'There is a smoothness and flow to the wing stroke
which is astonishing in so large a bird*. The flapping rate is
179/min (n = 13) (Kahl 1971b).
One of the world's largest storks, the Jabiru is consider-
ably larger than the American Wood Stork and has a mas-
sive upturned, rather than downturned, bill. Its size and
black head and bill distinguish it from the Maguari Stork. It
is a stately bird, that stands tall in the marsh, often alone or
in pairs. In flight, it is similar to an American White Pelican
(Pelecanus erythrorhynchos), but is larger, with extended neck
and legs and all white wings.
DISTRIBUTION AND POPULATION
The Jabiru Stork breeds from southern Mexico (Campeche,
Tabasco, possibly Quintana Roo), locally through the low-
lands of Central America (Belize, Guatemala, Nicaragua,
Costa Rica) and most of tropical South America east of the
Andes. Its overall South American range is from Colombia
(lower Magdalena), Venezuela and the Guianas, southward
through eastern Peru (rare on the coast), eastern Bolivia,
Brazil, Paraguay and Uruguay to northern Argentina (Cor-
rientes, Santa Fe Provinces; accidental to Buenos Aires)
Jabiru Stork 121
(Meyer de Schauensee 1970, Blake 1977:189-190, Koepcke
1963, pers. obs. J.A.K.). Populations are concentrated in
the vast wetlands of the Colombian and Venezuelan Llanos,
the Brazilian Pantanal (Mato Grosso) and the Paraguayan-
Argentine Chaco. Vagrants have been recorded north to
Texas and Oklahoma (Haucke and Kiel 1973, Oberholser
and Kincaid 1974:123, K.A. Arnold 1978).
In South America, the Jabiru Stork is generally resident
and non-migratory (B.T. Thomas 1985), Reportedly, it
leaves Belize between June and November (Luthin 1984a).
It is said to leave the main rivers in the Amazon region of
eastern Peru during the high-water season and to move to
smaller streams and higher country (Koepcke 1963:403).
Formerly, it was found in Honduras only in December and
January (Monroe 1968); recent reports are lacking. It does
wander, being recorded rarely as far north as Texas and
Oklahoma, as far west as coastal Peru, and as far south as
Buenos Aires, Argentina (Olrog 1963, Hughes 1970,
Haucke and Kiel 1973, K.A. Arnold 1978, C.T. Clark
1982).
Populations in Central America have historically been
small and highly localized. In recent surveys, only a few
nests have been documented in most areas of the region, and
the entire Central American population is estimated at
between 150-250 individuals. A maximum of 83 adults and
6 juveniles were counted along the Gulf coast of Mexico in
May 1978 (Knoder et al. 1980). Populations in northern
South America are larger, but nevertheless relatively low.
Only about 200 individuals were found on surveys in the
Llanos of Venezuela in 1983 (Ramo and Busto 1984),
although their dispersed nesting makes surveys difficult.
Coastal populations are generally smaller; a maximum of
100 birds was seen along the Surinam coast (Spaans 1976a).
The largest populations are found in the Pantanal and
eastern Chaco in south-central South America. Brooks
(1991) counted one bird per 4.38 km2 in the Paraguayan
Chaco. It is said to be very rare in Uruguay (Cuello and
Gerzenstein 1962:35).
ECOLOGY
Jabiru Storks favour extensive shallow marshes, wet mea-
dows, ponds, rivers and pastures, both inland and along the
coast. They are birds of open country but prefer wooded
areas nearby for roosting and nesting.
This species feeds by Standing or by Walking Slowly,
with the neck extended forward and the bill pointed down-
ward, while non-visually Probing every few steps. It will
also Stab at food items visually, Probe quickly, or Run and
Hop short distances after prey. When a prey item is cap-
tured, the birds frequently carry it ashore, bite it several
times before swallowing. Buried eels are sought by Probing
into the mud. In Venezuela, 43 individuals located about
half of their prey by feel and half visually (B.T. Thomas
1985).
Jabirus usually feed singly or in pairs, but when prey
becomes concentrated, such as by falling water levels, they
join loose, sometimes large, flocks, and mixed-species aggre-
gations. They also are attracted to fires. Large numbers
122 Jabiru Stork
were observed gathered during a mouse plague in Argen-
tina. 'Armies of these majestic white birds were seen stalk-
ing over the grass on all sides, or at the close of day winging
their flight to the distant watercourses in a continuous flock'
(Hudson 1951:58-59). Naumburg (1930:89) observed 200-
300 together at ponds in the Pantanal of the Mato Grosso,
Brazil.
Jabirus are resident in large territories, which they defend
(B.T. Thomas 1981). Males are highly aggressive in capti-
vity both to conspecifics and to others, including humans
(Shannon 1987). In Venezuela, a Jabiru aerially chased an
American Wood Stork that had caught a 30cm eel; how-
ever, the owner retained and managed to swallow the prey
(B.T. Thomas 1985).
The food is primarily fish and eels (e.g. Symbranchus mar-
moratus in Venezuela), but they can take many kinds of prey
including insects, snails (Ampullaria guianensis and A. papyr-
acca, in Guyana), mussels, crabs, frogs, small mammals and
snakes (Schomburgk 1922-23:121, Pelzeln 1868-71:305,
Chubb 1916:159, Meyer de Schauensee and Phelps 1978:22,
Ogden and Thomas 1985b, B.T. Thomas 1985).
BREEDING
The Jabiru Stork nests singly on the tops of tall broad-
leaved trees (e.g. Bombax munguba, Calycophyllum spruceanum,
Ceiba pentandra, Eriodendron anfractuosum, Enterolobium contorti-
siliquum), and often in palms (e.g. Copernicia tectorum, Butia
yatay). In coastal Mexico, nests are in tall red mangroves
(Rhizophora mangle). Nests are sometimes near mixed colon-
ies of other species but usually not near other Jabirus, which
are driven away from the nesting area. B.T. Thomas (1981)
found 5 active nests within a 250 ha area in Venezuela. The
large, stick nest is placed in or near the top of a large tree,
often one that towers over its neighbours and affords a view
of the surrounding country. Many authors have commented
on the Jabiru choosing the largest tree around for nesting
(Roosevelt 1914:90, Bent 1926:67-68, Naumburg 1930:90).
We have observed a typical situation in the species' north-
ernmost nesting site in the Usumacinta Delta, Mexico. The
nest was situated in a tall mangrove tree adjacent to man-
grove swamp and open marshes. The tree projected well
above the surrounding canopy with a commanding view of
the countryside. On rare occasions (Schomburgk, in Bent
1926:67) the birds may nest atop large rocks.
Nests themselves are flat, massive structures, up to 2 m
across and 1 m deep, made of branches up to 5 cm thick and
1.5-2.0 m long, intermeshed with smaller twigs and lined
with dry grass. Nests are often reused in successive years;
however, nests in palms are not reused if the palm dies. A
new nest in Venezuela took nearly 2 months to complete
(Hagmann 1907, Kahl 1971e, B.T. Thomas 1981).
The nesting season is variable across the range (see
Appendix) and seems largely determined by seasonal rain-
fall patterns. In both Venezuela and Argentina, birds nest
from the latter part of the rainy season into the dry season
(Kahl 1971e, B.T. Thomas 1981). In Surinam, they nest in
the long dry season (Spaans 1975b).
Especially during the nesting season, Jabirus may exhibit
intra- and interspecific territoriality near the nest and at
feeding areas. B.T. Thomas (1985:928) recorded that
Jabirus twice chased Maguari Stork fledglings that flew
within 100m of Jabiru nest sites. A captive male has been
observed, threatening a tapir by throwing his head back
until the bill was vertical and snapping the bill, and males
were very aggressive toward their keepers in the breeding
season (Shannon 1987:245).
It is likely that solitary pairs remain mated in successive
seasons and use the same nesting site, so courtship display is
not extensive (Kahl 1971e, 1973b). On the feeding grounds,
pairs engage in Flap-Dash displays, which may function in
pair-bond maintenance. In captive birds (Shannon 1987),
pair-formation is initiated by the female, who overcomes the
aggressive territoriality of the male by using a display in
which she rushes up to the male with her head down while
coughing and bill clattering. The first greeting displays also
appear to be initiated by the female, with the male respond-
ing.
After pairs have formed, courtship displays are those
typical of storks (Kahl 1973b). The Up-Down display in
greeting involves the birds facing each other, with bodies
erect, wings closed, necks upward and bill nearly horizontal.
The birds then bill-clatter loudly and rapidly (8-10 times/
s), while slowly swaying their heads and necks from side to
side and slightly raising and lowering the bill across the
horizontal plane.
The Flap-Dash is a spectacular social display, usually
seen away from the nest at feeding areas. A member of a
pair, usually the male, suddenly stands erect, runs away
from its mate with open and flapping wings, then turns and
dashes back to the mate, and stands for a moment with
wings fully spread (Kahl 1973b). This display has been
reported only in Jabiru, and in the Blacknecked and Saddle-
bill storks (genus Ephippiorhynchus), the only three species of
stork with white flight feathers in the wing. In wild Jabiru
Storks this display was usually seen performed by pairs, but
Shannon (1987) saw it performed in captivity by single
birds as well.
Arching displays, with loud bill snaps, occur in response
to disturbance near the nest. Copulation takes place on the
nest, usually begins without preliminary display and lasts
13-19 s; Copulation Clattering may involve gasping coughs
from one of the birds (Kahl 1973b, B.T. Thomas 1981).
During early nest-building, the sticks are mainly gathered
by the male and positioned by both birds (Kahl 1971e,
Shannon 1987).
Both sexes incubate and feed the young. When disturbed
at the nest, adults give loud claps of their bill. (Lloyd 1895,
Schomburgk 1922-23:121).
The eggs are a dirty white, laid in clutches of 2-4, usually
4 but sometimes 5 (Lloyd 1895; Hagmann 1906; Kahl
1971e). 11 nests studied by B.T. Thomas (1981) in Vene-
zuela contained the following: single chicks (2 nests), 2
chicks (5 nests), 3 chicks (2 nests), 4 chicks (1 nest), or 5
chicks (1 nest). The fledgling period was estimated at 10-11
weeks, but this is based upon observations at a single nest
and may be something of an underestimate (B.T. Thomas
1981, 1985).

TAXONOMY
The Jabiru Stork shares many characteristics with the Sad-
dlebill and Blacknecked Storks (genus Ephippiorhynchus] on
the one hand, and with the Lesser and Greater Adjutant
and Marabou Storks (genus Leptoptilos) on the other (Dek-
eyser 1962, Kahl 1971e). The Jabiru appears closer beha-
viourally to the former and morphologically to the latter.
We retain its monotypic genus to demonstrate this inter-
mediate taxonomic position (Kahl 1972e).
CONSERVATION
Habitat destruction and human disturbance is recognized
as a primary threat to the Jabiru throughout its range
(Luthin 1987). Wetland drainage, especially for agricultural
conversion, tree cutting and hunting are the principal
threatening human activities.
Jabiru Stork 123
The Jabiru has become a symbol of conservation and
wetland protection in Central America, particularly in
Belize and Costa Rica. It is considered to be threatened in
Central America and endangered in Nicaragua (Luthin
1984a). Specific needs include population ecology studies
and the creation of Jabiru reserves to protect wetland habi-
tats important to many species (Luthin 1984a).
In Surinam, the species is 'uncommon nowadays and
reduced by relentless shooting' (Haverschmidt 1968:28). It
is, in fact, considered to be a gamebird in the Amazon, and
larger nestlings are particularly sought (Sick 1985).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 301.
 Lesser
Adjutant Stork
Leptoptilos javanicus (Horsfield)

Ciconia Javanica Horsfield, 1821, Trans. Linn. Soc. London, 13, p. 188: Java

Other names:
Hair-crested Adjutant, Smaller Adjutant, Javan Adjutant, Hair-crested Stork, Marabou Crane
(English); Marabout d'Asie (French); Javaanse Maraboe (Dutch); Malayischer Kropfstorch,
Stirnplatten-Marabu, Sunda-Marabu (German); Chinjara, Chandana, Chandiari, Bang gor, Chota
garur (Hindi); Madan-chur, Madantak, Modun-tiki (Bengali); Tokla moora [='bald head'], Bor
tokola (Assamese); Dodal kongo, Dodal gatti gadu (Telugu); Meva kokku, Mana (Tamil); Mana
kokka (Sinhala); Vayalnaiicken (Malayalam); Don-mi-gwet (Burmese); Tontsap (Arakan); Bangau,
Bango (Javanese); Burong botak [='with bald head'], Burong babi (Malay); Burong Kambing,
Burong Gajah (Sumatran); Hoang (Hainan Chinese)

IDENTIFICATION
The Lesser Adjutant Stork is a massive, heavy-bodied stork, standing 110—120 cm tall, if erect, but often
shorter because it stands hunched with the head drawn in on the shoulders. It has a dark grey back,
white underparts, and sparsely feathered head and neck. It is generally dignified and sluggish in
demeanour.
This stork is glossy metallic blue-black above, white below, with sparse hairlike feathers on the nearly
naked, reddish yellow head and neck. The bill is dull yellow, with the tip whitish and the base tinged
red to pinkish. The bare crown or 'frontal plate' varies from greenish brown to light yellowish tan. The
featherless face and neck are a mixture of pale pink and bright tumeric-yellow, tinged with brick-red.
The iris is pale slaty grey to pale bluish grey. The legs and feet are greenish brown to almost black but
are usually difficult to see because they are covered with white excreta.
The sexes have identical plumages. However, the males are noticeably taller and heavier, with a more
massive bill. This is a distinction that can be recognized in wild birds.
At the start of breeding, the back remains a dark slaty blue-black, and small, oval-shaped copper
spots about 2.0 cm long develop near the tips of the median secondary coverts (these spots are not on
the greater coverts, as sometimes stated in the literature). Narrow white edging appears on the
scapulars and 4-5 of the innermost greater secondary coverts. The other greater secondary coverts
remain dark.
Nestlings are covered with white down, except for the head and face, which are largely naked. The
juvenile is similar to the adult, except that the upperparts are less glossy, and they lack the copper spots
on the coverts. The head and neck are more densely feathered than in the adult.
During the Up-Down display, given at the nest, birds utter a series of high-pitched squeaks and
cowlike 'Moos'. Vocalizations heard during these displays were hoarser and more rasping than those
given by Greater Adjutant Storks. Their quality was reminiscent of some of the vocalizations of young
Marabou Storks at about the time of fledging (Kahl 1970, 1972a). M.C. Thompson (1966) reported
calls resembling the 'deep growl of a dog' from non-breeding birds in Borneo. A loud 'Woofing' wing-
noise can be heard near the nest.
126 Lesser Adjutant Stork
Unlike other storks that fly with the neck outstretched,
birds in the genus Leptoptilos fly with the head drawn back
on their shoulders (Glenister 1951, Henry 1971, Kahl
1972a). The Lesser Adjutant commonly soars when thermal
air-currents are available; soaring/gliding flight is a
favoured mode of transport between nesting/roosting areas
and foraging grounds. The flapping rate has been timed at
153/min(w = 8)(Kahll971b).
The Lesser Adjutant can be most easily confused with the
Greater Adjutant. It is distinguished by its smaller size;
somewhat thinner and slightly recurved bill; dark slate-
black (as opposed to paler bluish grey) back and upperside
to wings; pale light-tan forehead; more dense down and
hairlike feathering on the neck and back of head, most
pronounced in a small crest on the occiput; much smaller
(rarely even noticeable) gular pouch, which is actually only
a band of expandable, bare, reddish skin around the base of
the neck and is not pendant, feathers that form a white ruff
at the dorsal base of the neck with large black spots; white
(not sooty grey) undertail-coverts; tibia black up to the
feathers. These differences are illustrated here, and in colour
in plate VIII of Kahl (1971f). The adult Lesser Adjutant
resembles the Juvenile Greater Adjutant in many aspects of
morphology.
DISTRIBUTION AND POPULATION
The Lesser Adjutant Stork occurs in tropical Asia and
adjacent islands. It was once widespread from India
(mainly eastern and southern) and Sri Lanka to southern
China, Vietnam, Thailand, Malaysia, Sumatra, Java and
Borneo. As far as is known, it is only an occasional visitor to
Nepal and Bali (M.D. Bruce 1982) and apparently no
longer breeds in Burma or Thailand.
This is mainly a non-migratory species. However, it can
be nomadic, especially in response to rains. It may be
locally migratory, although this needs confirmation. The
Borneo population is reported to increase in the winter, with
up to 200-300 birds seen there in January (Smythies
1981:31).
There are few censuses available. Formerly this species
was less abundant than the Greater Adjutant (E.C.S. Baker
1929:330); today, however, the Lesser is the more common
species in almost all areas of their common range. The
population in India is now more thinly scattered, with
recent reports of nesting only in Madhya Pradesh, Assam
and perhaps the Indian Sundarbans (Luthin 1987, P. Saikia
and P.C. Bhattacharjee pers. comm. 1990).
During a survey in Assam, India, during 1987-89, it was
estimated that there were about 400 Lesser Adjutants. The
survey found 40 nests in Orang, 11 in Manas, 20 in Kazir-
anga, 11 in Lowkhowa reserves, and 53 nests outside pro-
tected areas (Saikia and Bhattacharjee 1989a,b).
Small breeding colonies in Yala National Park, Sri
Lanka, number about 100 pairs. A single colony is known
from Vietnam, and no information is available from Laos
and Kampuchea (Luthin 1987).
A population of about 200 birds is estimated to occur on
the west coast of peninsular Malaysia (D. Wells pers.
comm. 1989). In Sumatra, its main stronghold is in South
Province. As many as 620 Lesser Adjutants were counted
there during 1984—89. The entire Indonesian population
(Sumatra, Java, Kalimantan, Bali) is believed to be no
more than 2000 (M.J. Silvius and WJ.M. Verheugt pers.
comm. 1990). Elsewhere no data exist on population
numbers.
ECOLOGY
The details of the Lesser Adjutant Stork's ecology are little
known. It inhabits flood-plains, swamps, pools in forests,
marshes, drying river beds, lakes, and rice-paddies. It some-
times moves to agricultural fields and grasslands. Particu-
larly in Southeast Asia and the Greater Sunda Islands, it is
a bird of mangroves and coastal mudflats.
The Lesser Adjutant is apparently less of a scavenger
than the Greater Adjutant or Marabou Storks. It feeds
alone or in small flocks and shows considerable alertness
and activity when hunting for food. Most food location
appears to be visual, with the bird Walking Slowly on dry
ground or shallow water and Grabbing prey in the tip of its
bill. We have seen up to 6 birds perched on top of wooden
telephone poles lined across a marsh in Assam, apparently
scanning the marsh from their high vantage point for poten-
tial prey (J.A.H. pers. observ.)
Food consumed includes fish, frogs, reptiles, crustaceans,
locusts, rodents, small mammals, lizards, insects and some
carrion (Kahl 1970). We have seen birds bring whole ver-
tebrates, such as frogs, fish and rats, back to the nest. The
Lesser Adjutant is in fact known to 'eat any living thing,
including chickens, not too big to swallow' (E.C.S. Baker
1929:330).
Because of its preference for natural prey, the Lesser
Adjutant is less attracted to the vicinity of human habitation
than is the Greater Adjutant. While resting, it often sits on
its intertarsal joints, with the tarsi stretched out in front.
BREEDING
In India and Burma, nests are placed from 12-30m up in
tall forest trees, such as satian (Aistronia scholaris) or simul

(Salmalia malabarica) (Kahl 1970). In Southeast Asia, it
breeds in coastal mangroves on the shoreward edge of man-
grove forests or along the banks of tidal rivers (H.C. Robin-
son and F.N. Ghasen, 1936:210). Some tree-nests in
southern Burma were as high as 46m (Hopwood 1921,
E.C.S. Baker 1935:448).
The Lesser Adjutant Stork usually nests near water-
bodies, although the water may dry before breeding is
complete (P. Saikia and P.G. Bhattacharjee pers. comm.
1990). It is a semi-colonial or colonial breeder. It generally
breeds in small colonies, sometimes with as many as 12-20
nests in a single large tree (E.C.S. Baker 1935:448). In the
past, at least, it bred in mixed colonies with the Greater
Adjutant (Smythies 1953:523).
The nesting season of the Lesser Adjutant in Assam,
India, can begin as early as July (P. Saikia and P.C. Bhatta-
charjee pers. comm. 1990). In southern Burma, where they
formerly nested in large numbers in the company of the
Greater Adjutant, the Lesser Adjutant was said to begin
nesting somewhat later than its larger cousin (Blanford
1898:375). However, such was not the case in Assam, India,
where we observed nests of both species in November-
December 1967. Here 36 nests of the Lesser Adjutant con-
tained young 35-75 days old, whereas a week later a single
nest of the Greater Adjutant was found with young about
30-35 days old (Kahl 1972a).
The nest is a large but rather flimsy stick structure 60-
150 cm in diameter and 10-120 cm deep, lined with smaller
twigs and green leaves. Old nests from previous years are
often used again by birds who add sticks to the old foun-
dations in the new season.
The general courtship behaviour of the Lesser Adjutant
appears to be very similar to that of the Marabou Stork in
Africa, with the exception of a few specific displays.
The Balancing Posture is given by females during pair-
formation. This display in the Lesser Adjutant involves
lifting the head in a scooping motion and contains a bill-
clattering component lacking in both the Greater Adjutant
and Marabou (Kahl 1972a). During the Up-Down display,
given as a greeting between mates at the nest, both species
of Adjutant clatter while the bill is held above the horizon-
tal, whereas the Marabou clatters with the bill below the
horizontal.
We have observed few threat displays in this species. The
Bill Snap, Pre-flight Snap and Erect Gape are similar to
those of the Greater Adjutant and Marabou. In the Arching
display, given in the presence of intruders, the Lesser Adju-
tant extends its neck and gives a hoarse wail. The Arching
displays that we saw, were not as ritualized as in the
Marabou (Kahl 1972a).
Eggs are white, but become quickly soiled during incu-
bation. A full clutch contains 2-4 eggs. The eggs of this
species are quite similar to those of the Greater Adjutant,
and in spite of the considerable size-difference of the
two species, their eggs differ only slightly (E.C.S. Baker
1935:449). The incubation period is 28-30 days (P. Saikia
pers. comm. 1990).
Both adults feed the nestlings; young are also given water
in the nest by their parents. Some observers have reported'
Lesser Adjutant Stork 127
the Lesser Adjutant to be more shy than the Greater; how-
ever, this was not found to be the case at the nest (Kahl
1-970). Young often gave the Up-Down display to their
parents before the Begging display (Kahl 1972a).
In an odd situation, at the Dehiwela Zoo, Columbo, Sri
Lanka, a female Lesser Adjutant hybridized with a male
Painted Stork (Mycteria leucocephald). Both parents were free-
flying and ranged over neighbouring paddyfields at will.
The 'odd couple' hatched 4 broods between 1940 and 1942,
and a number of viable offspring were produced. One could
still be seen walking freely about the zoo as late as
December 1967 (pers. obs. M.P.K.). The hybrid offspring in
adulthood most resembled the Lesser Adjutant Stork in
plumage-coloration and bill-size, and the Painted Stork in
stance and bill shape (W.C.O. Hill 1943, Henry 1955, A.P.
Gray 1958, pers. obs. M.P.K.). The same species combi-
nation (sexes unknown) also hybridized in 1974 at the
Kuala Lumpur Zoo, Malaysia, and produced young (P.
Olney pers. comm. 1974).
TAXONOMY
Of the three species in the genus Leptoptilos, the Lesser
Adjutant is the least specialized in its morphology and
behaviour. In some morphological characters, such as bill-
shape and external structure of the gular pouch, the Lesser
Adjutant shows more affinities with the Jabiru than do the
other two Leptoptilos species (Kahl 1971e).
CONSERVATION
Numbers of this stork appear to be much reduced over most
of its range. In Assam, India, habitat loss, both from the
cutting of nest-trees and the draining of feeding areas, is a
threat to the Lesser Adjutant. At least one ethnic group kills
the birds so that the bill and skull can be used for medicinal
purposes (Saikia and Bhattacharjee 1989a,b).
This species is poached in Bangladesh and is sold in the
market in Bangkok, Thailand (M.A.R. Khan 1987, Luthin
1987). Although there are about 200 birds in peninsular
Malaysia, there is very little breeding there owing to the loss
of suitable large mangrove trees and the robbing of young
by humans. Birds are still widely but thinly distributed in
Sabah, Borneo, especially on the remote northeastern and
eastern coasts (D. Wells pers. comm. 1989). However, they
are said to be decreasing rapidly there, probably owing to
shooting (Gore 1968).
As far as we can determine, the large numbers that once
bred in southern Burma no longer exist. The healthiest
population may be in southeastern Sumatra, but the birds
there are at risk because of swamp-forest clearance, direct
persecution and transmigration of people into the area (M.
Silvius and W. Verheugt pers. comm. 1990).
A few breeding sites are protected (in Assam, the Sundar-
bans, and Vietnam), but even there human disturbance and
poaching continue.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 302.
 Greater
Adjutant Stork
Leptoptilos dubius (Gmelin)

Ardea dubia Gmelin, 1789, Syst. Nat., 1, p. 246; based on 'Gigantic Crane' of Latham, 1785, General
Synop. Birds, 3, p. 45: India

Other names:
Adjutant Stork, Larger Adjutant, Pouched Adjutant (English); Indische Marabu, Argala, Argala-
Marabu (German); Hargila [=bone-swallower], Garur, Peda dhauk (Hindi); Dusta (Dakhani),
Dhenk (Mirshikars, Bihar); Hargila, Chaniari Dhauk (Bengali); Hargila, Bortukla (Assamese);
Peenigala konga (Telugu); Don-zat (Burmese)

IDENTIFICATION
The Greater Adjutant Stork is the most massive and ugliest of the Asian storks, standing 120-150 cm to
the top of the head when erect. It often appears shorter, owing to its hunched stance. Its general colour
is slaty-black, grey and white, with a nearly featherless head and neck, a massive wedge-shaped bill and
a naked ruddy pinkish gular pouch, hanging from the front of the neck. It was named 'Adjutant'
because it walks with the deliberate, measured gait of a military adjutant.
The adult's upperparts are slate grey, with a pale band along the wing formed by the pale grey
greater secondary coverts. The underparts are white. The head, neck and gular pouch are almost
naked, with a few scattered dark brown hairlike feathers on the nape, neck, and sides of head. Dorsally
there is a ruff of white feathers around the base of the neck, surrounding a red or orange air sac, c. 15 cm
in diameter, that is only visible when inflated.
The iris is white to yellowish white. The bill varies from pale yellowish to greenish, more reddish near
the base. The bare skin of the head is a dull reddish brown, turning to brick-red on the hindneck and
blackish on the forecrown. The gular pouch and neck are mostly yellow, with more pink, spotted with
black, on the pouch. The legs and feet are actually a horny brown to dark grey, but often appear pale
greyish white or chalky white because they are coated with uric acid from excreta.
The male and female have the same plumage and soft-part colours. Sexual dimorphism in size is
slight. However, males are somewhat larger than females, with heavier bills; the female's bill is a bit
more decurved.
At the start of breeding, the upperparts become a pale bluish grey, and the paler wing band turns a
silvery grey. The undertail-coverts are white at their bases and dark smoky grey at their tips; they become
luxuriant and fluffy at breeding time. The face and forehead become encrusted with what appears to be
dried blood. The gular sac becomes a bright saffron-yellow (see colour plate VIII in Kahl 1971f).
Nestlings are covered with a dense white down, except for the bare head and face. They are similar in
appearance to nestlings of the Lesser Adjutant and the Marabou, except that Lesser Adjutant nestlings
have a slightly thinner bill.
Juveniles are similar to adults but have darker uppersides, without the pale band along the top of the
wing, and with denser pale-brownish and grey feathering on the head and neck. Like adults, they also
have smoky grey undertail-coverts.
130 Greater Adjutant Stork
This is generally a silent bird, except around the nest.
Bill-clattering is common during various displays. During
displays at the nest a high-pitched squealing, slurring to a
lower-pitched cow-like 'Mooing', is uttered; these calls are
quite similar to those given by the Marabou Stork of Africa
(Hume and Gates 1890:262, Kahl 1966a, 1972a).
When the wings of adults are viewed from above in flight,
there is a conspicuous pale grey band formed by the silvery
grey greater secondary coverts. Despite heavy flapping on
take-off, the bird flies strongly with noisy rhythmic flaps
when once airborne and circles gracefully in thermals. 'On
the wing .. . they are very majestic and a flight of these birds
sailing round in great circles is very imposing' (E.C.S. Baker
1929:329). Unlike other storks, birds in the genus Leptoptilos
fly with their necks partially retracted back onto their
shoulders, similar to the herons. The flapping rate has been
timed as 139/min (a = 8) (Kahl 1971b).
The only other species within its range with which the
Greater Adjutant could be confused is the Lesser Adjutant.
The Greater is distinguished by its larger size, paler grey
upperparts, pale grey band along the upperside of wing
(which is conspicuous both at rest and in flight), ruff of
unspotted white feathers around the dorsal base of neck,
more massive bill (often with a slightly decurved culmen),
dark (as opposed to pale-tan) forehead; dark sooty grey (not
white) undertail-coverts, more sparsely feathered head and
neck, c. 50 mm of red skin visible at the upper end of
unfeathered tibia; and much larger gular pouch (20-35 cm
long when inflated). All these differences are shown in plate
VIII of Kahl (1971*).
DISTRIBUTION AND POPULATION
The Greater Adjutant is sparsely distributed in tropical
Asia. Formerly it bred from northeastern India, Burma to
Thailand, Kampuchea and southern Vietnam; it has now
been extirpated or is uncommon in most of its former haunts
(Deignan 1963:10, Wildash 1968:46, B.F. King and E.G.
Dickinson 1975:52, S. Ali and S. D. Ripley 1968:106).
Recent breeding is known only in Assam, India (Kahl
1970, Saikia and Bhattacharjee 1990b, A. Rahmani pers.
comm. 1990). However, the species may still be nesting in
the swamp forest of Minh Hai Province, Vietnam, although
it has not been sighted in recent years (Luthin 1987).
Earlier accounts of the range extending to the Greater
Sunda Islands (Sumatra, Java, Borneo) (E.C.S. Baker
1929:328, J.L. Peters 1931:130, S. Ali and S. D. Ripley
1968:106) are almost certainly in error, being instead the
Lesser Adjutant (Kuroda 1936:542). According to David
Wells (pers. comm. 1989) there are 'no acceptable records
[of Greater Adjutants] south of Thailand/
The Greater Adjutant is largely resident but also noma-
dic, depending on local conditions. During the period of
their former abundance, Greater Adjutants came to India
mainly during the rains and then returned to Burma to
breed during the dry season. In years when desert locust
swarms were heavy, the storks sometimes appeared in large
numbers in areas of India (e.g. Rajasthan) where they
usually were not common (Singh and Singh, 1960). Birds
wandered irregularly, primarily during the rains, to Pakis-
tan, Nepal, north-central India, and south (rarely) to Tamil
Nadu ( — Madras Province).
In the last few years, few, if any, Greater Adjutants have
been reported outside Assam (none at Bharatpur since
1987; Bholo pers. comm. 1990), and this may suggest that
nearly all the original migratory stock which formerly bred
in Burma have been extirpated, and that the isolated popu-
lations in Assam are mainly sedentary as reported by P.
Saikia and P.C. Bhattacharjee (pers. comm. 1990).
Up through the 1930s very large numbers ('literally in
hundreds of thousands', according to E.C.S. Baker
1929:328) gathered to breed in Burma, and it was thought
that perhaps all of the Indian population of Greater Adju-
tants went there each year at the end of the rains (Hume
and Oates 1890:261). The immense breeding colonies once
reported near Shwaygheen, in the Pegu District of southern
Burma (c. 150km northeast of Rangoon) are apparently
gone today. Similarly the colony site in the Sunderbans of
Bangladesh has been eliminated (M.A.R. Khan 1987).
The species was formerly 'rather common' in Chiang Rai
Province, northern Thailand (Deignan 1945:46), but it is
much reduced in that country now (Lekagul 1968). Even as
far back as the 1920s it was apparently much less common
in Indochina than was the Lesser Adjutant (Delacour and
Jabouillc 1931:89).
Between 1987 and 1989 the largest number of Greater
Adjutants seen in Assam, India, on a single day was 56 in
Tezpur and 83 in Guwahati, and the total population there
was estimated at 300 birds (Saikia and Bhattacharjee
1989a,b). In April-May 1989 Rahmani (1989c) counted 80
Greater Adjutants at 10 locations in Assam. More recently,
in October 1989-January 1990, a special search was
launched, and 75 active nests were located in Lower Assam
(Saikia and Bhattacharjee 1990b).
ECOLOGY
Greater Adjutant Storks occur in various wetlands, such as
jheels, marshes, paddyfields and lakes, particularly 'where

the water is drying and concentrating the fish life in shallow
puddles' (S. Ali and S.D. Ripley 1968:106). Their use of
wetland habitats is primarily during the breeding season;
during the non-breeding period they often feed on human
refuse or at abattoirs.
Formerly the species was very common around human
habitations, where it fed on carrion. During the 19th Cen-
tury, 'large numbers of [Greater] Adjutants haunted the
city of Calcutta . . . for the offal and refuse, and rendered
valuable service as scavengers' (S. Ali and S.D. Ripley
1968:106). 'At that time . . . Adjutants could be seen on the
highest points of almost every house in Calcutta' (E.C.S.
Baker 1929:329).
The usual feeding behaviour is for the bird to Walk
looking for food, or alight at a food source and Stand to eat
what is available. Food finding is primarily visual.
It has been suggested (Jerdon 1864:730) that the inflated
gular sac rriay help to supply extra oxygen 'during pro-
tracted acts of deglutition'. We doubt this. We feel that the
gular sac functions primarily during social displays and,
possibly, in thermoregulation.
When feeding on swarms of the desert locust in Rajas-
than, India, a stork was reported running a few steps and
'flapping its wings briskly to beat the bushes . . . on which
large concentrations of the hoppers and fledglings of the
desert locust were noted to be resting in the morning.
Disturbed by the heavy and noisy flaps of the wings the
insects jumped down from their roosting places, when they
were immediately attacked with the wedge-shaped bill,
picked up, and devoured' (Singh and Singh 1960).
The food includes carrion, fish, frogs, reptiles (Vipera
russelli, Uromastix hardwickii), crustaceans, locusts, and even
maimed ducks (Panday 1974). It is likely that the diet
changes during the breeding season to include many whole
vertebrates for the essential calcium requirements of the
fast-growing young.
Outside the nesting season, this is a species well known
for its appalling eating habits. According to E.C.S. Baker
(1929:329) 'their tastes are omnivorous and there are few
things an Adjutant will not swallow'. According to Akhtar
(1947) a captive Greater Adjutant 'swallowed a shoe well-
shod with iron; on another occasion it swallowed a good-
sized fowl right down, with its wings and feathers.' In
Assam, India, one bird was seen swallowing pieces of ver-
tebral column from a dead buffalo that were over 30 cm long
(Rao and Murlidharan 1989). A local Indian name, 'har-
gila', means 'bone-swallower'.
The birds are fairly gregarious in foraging, associating
with other Adjutants and with vultures and kites at animal
carcasses and refuse dumps. When using wetland sites, they
feed until the food supply is exhausted; then the group
moves on to another site (P. Saikia and P.C. Bhattacharjee
pers. comm. 1990). We have seen aggressive encounters of
captive birds at feeding sites elicit Forward Displays and
Upright Displays (Kahl 1972a). But even adult Adjutants
are often dominated by the more aggressive vultures at
feeding sites. The Greater Adjutant is often found in and
around urban centres, where there are sources of food, and
so seems not disturbed by close association with man.
Greater Adjutant Stork 131
BREEDING
The Greater Adjutant Stork nests, as well as roosts, in tall
trees, often near villages and not necessarily in wetlands. In
Burma it has nested on limestone pinnacles. A nest we saw
in Assam, India, was placed in the crown of a satian tree
(Aistronia scholaris) about 27-30 m from the ground; this nest
was in the tallest tree within 500m (Kahl 1970). In late
1989, 75 nests were found concentrated in 6 colonies in
Assam, India. They were placed at heights of 12-23 m in a
variety of trees, such as kadam (Anthocephalus cadamba),
simul (Bombax ceiba) and others. Nest trees were surrounded
and protected by dense stands of bamboo (Saikia and Bhat-
tacharjee 1990b). The nest itself is a large platform of coarse
sticks, 1-2 m in diameter and up to 1 m deep.
Pairs may breed singly, semi-colonially or colonially,
sometimes with other birds, including the Lesser Adjutant
Stork. At recent Assamese sites, some pairs nested singly,
and the greatest number of nests in a single colony was 31
(Saikia and Bhattacharjee 1990b).
Eggs are layed from September to January in Assam. In
some areas, this species has been said to begin breeding
about a month earlier than the Lesser Adjutant (Hume and
Oates 1890:260). However, that was not the case with a
single nest found at Kaziranga Wildlife Sanctuary, Assam,
India, in late 1967. Nor was it the case in Assam in 1990,
where some nests were still being built in November, after
most Lesser Adjutants had fully grown young (pers. obs.
J.A.H.).
Most courtship displays described for the Greater Adju-
tant (Kahl 1972a) are similar to those shown by the Mara-
bou Stork of Africa, which has been studied in more detail
(Kahl 1966a).
Adults are quite noisy at the nest, with many displays
incorporating a loud clattering of the bill, plus a variety of
squealing and 'Mooing' calls and loud wing noise. The Bill
Snap is used most commonly when birds were being
approached by another adult: the bird holds its body either
erect or horizontal, with neck retracted, and gives a single,
audible snap with the bill. It also gives a pop in the Pre-
flight Snap. The Erect Gape is given without vocalizations.
The Arching display is given in response to disturbance,
accompanied by a repeated high-pitched wail. We did not
see this display in the fully ritualized form, with neck puffed
out and wings fully spread, as occurs in the Marabou Stork.
The Balancing Posture is given, mainly by courting females,
in submission; Swaying Twig-Grasping is given mainly by
unmated males, as they are approached by a female.
The Up-Down display, given as a 'greeting' between
mates at the nest, terminates with the bill being clattered
upward in this species, instead of the downward clattering
seen in the Marabou.
The clutch size is 2 or 3 eggs, rarely 4; at 75 Assamese
nests in 1989, the average was 2.9 eggs (Saikia and Bhatta-
charjee 1990b). The shells are white, usually becoming
badly soiled as incubation advances. Both sexes share in
nest-building, incubation (E.C.S. Baker 1935:448) and feed-
ing of the young. The incubation period is 28-30 days (P.
Saikia pers. comm. 1990). At 75 Assamese nests in 1989-90,
132 Greater Adjutant Stork
c. 75% of the eggs laid produced fledged young, giving an
average of 2.2 per nest (Saikia and Bhattacharjee 1990b).
Young often give Up-Down displays to their parents,
before the Begging display. They gape widely, throw their
head to vertical, and then give several nasal squawks, before
starting the repetitious movements of Begging.
TAXONOMY
In many aspects of behaviour and morphology, the Greater
Adjutant closely resembles the Marabou Stork of Africa.
Some authors (e.g. Meinertzhagen 1951) have even sugges-
ted that the two species are closely enough related to be
considered conspecific. However, studies of both species at
the nest (Kahl 1966a, 1972a) have shown that one of their
principal courtship displays, the Up-Down, differs in sig-
nificant ways, and this difference would probably be suf-
ficient to provide reproductive isolation if the populations
were sympatric. Therefore, we feel it is biologically sound to
consider L. dubius and L. crumeniferus as valid, separate
species. However, they do appear to be closely enough
related to form a superspecies (Amadon 1966).
CONSERVATION
This is, perhaps, the most endangered of all the Asian
storks. In the early years of the 20th Century, the Greater
Adjutant was a very common bird in Indian cities during
the non-breeding season, and they gathered in immense
numbers to breed in southern Burma (Smythies 1953).
Today the bird is uncommon to rare in many of its former
haunts, and the Burmese breeding grounds are said to be
deserted (C. Luthin pers. comm. 1989). Investigations to
discover currently active breeding sites and measures to
protect them are urgently needed. If the decrease of the past
50 years continues at the same rate, the species might cease
to exist in another few years.
It is puzzling that the Marabou Stork seems to be increas-
ing in Africa, whereas the apparently ecologically similar
Greater Adjutant is decreasing in Asia. In India the species
has rarely been persecuted directly, for, being a carrion-
eater, it is considered by Hindus to be unclean and therefore
inedible. It is believed that many of the trees in the tradi-
tional Burmese nesting areas have been cut and the stork is
probably eliminated from that country (J. Sayer pers.
comm. 1989). The Greater Adjutant colony site in the
Sunderbans, Bangladesh, has been lost because of logging
operations going back into the 19th Century (M.A.R. Khan
1987); and the stork has been eliminated from elsewhere in
Bangladesh by hunting, except for one recent sighting in
December 1988 (A. Rahmani pers. comm. 1990).
The discovery, in late 1989, of 75 breeding pairs of
Greater Adjutants in Assam, India, is encouraging (Saikia
and Bhattacharjee 1990b). A major problem is that most of
the nest-trees are on private land, and the owners often cut
them down for their own use. Wetlands have been drained
for agriculture and remaining sites have been filling in, due
to the enormous growth of water hyacinths and grasses.
Industrialization is also encroaching on wetlands. Other
possible reasons for decline in India are pesticide poisoning
and increased competition from larger populations of vul-
tures (A. Rahmani pers. comm. 1990). Furthermore, adults
and nestlings are sometimes killed for food by members of a
nomadic (non-Hindu) tribe that visits the nesting areas
during the winter months, when the birds are breeding
(Saikia and Bhattacharjee 1990b).
The problems of the Greater Adjutant Stork are further
complicated by the fact that the local human population is
completely unaware of the status and need for protection of
this species. None the less, the Brahmaputra Valley of
Assam is the only site where conservation measures for this
species would have a chance of success. P. Saikia and P.C.
Bhattacharjee (1990b, pers. comm. 1990), who assessed
nesting locations, population status, and habitat use, pro-
vided a conservation strategy. This includes increasing the
conservation awareness of villagers, protection of nest-sites,
planting of roosting and nesting trees, and providing finan-
cial help to villages where nests are located. Known nest-
trees could be purchased as 'national treasures' and strict
laws against any harm to the birds should be enforced.
We strongly urge that the Indian Government take im-
mediate steps to ensure the safety of this rare species within
their borders.
Note'. Body measurements, egg measurements and nesting
season date for this species can be found in the Appendix on
page 303.
 Marabou Stork
Leptoptilos crumeniferus (Lesson)

Ciconia crumenifera Lesson, 1831, Traite Ornith., livr. 8, p. 585: Senegal

Other names:

Marabout d'Afrique (French); Afrika-Marabu (German); Abu Sen (Arabic); Bambu (Somali);
Jimmu (Mandingo); Borintunke (Hausa); Mmakaitsimeletsa (North Sotho); Svorenyama (Shona);
Qandlopfu (Tsonga)

IDENTIFICATION
This massive extremely ugly stork stands about 120cm tall, when erect rather than in its more usual
hunched posture, and has an impressive wing-span of up to about 2.9 m. It is basically slaty black, grey
and white, with a nearly featherless head and neck and a massive wedge-shaped bill. A distinctive
feature is its gular pouch of nearly naked ruddy pinkish to pale magenta skin. When inflated, the air
pouch hangs down 25-35 cm from the front of the neck.
To some eyes, the Marabou seems one of the ugliest birds in the world. It has been referred to as a
'gaunt-looking object at the best of times', 'the world's ugliest bird', 'entirely lacking any charm', 'stiff-
jointed and dried-up as though it had lived for a thousand years', or 'a dirty old man with a skin disease
of the scalp'. However, biologists who know the Marabou think otherwise and seldom speak of it in such
derogatory terms; we tend to notice the more endearing qualities in the bird.
Referring to the lovely white undertail-coverts, grown in the breeding season and formerly used in
high fashion, F J.Jackson (1938:79) stated: 'It is almost inconceivable that such delicate and beautiful
plumes, when adorning a fan or some finery dear to a woman's heart, are the product of this huge, gaunt
and repulsive looking scavenger, with a bald, scabby head and a body alive with vermin.'
The upperparts are dark slate-grey, the underparts white. The flight feathers are glossed with green,
whereas the back and mantle are glossed with blue. Head, neck and gular pouch are nearly naked, with
a few scattered dark reddish brown, hair-like feathers on the nape, neck, and sides of head. Dorsally, at
the base of the neck, there is a ruff of white feathers, surrounding a deep crimson or orange-red air-sac c.
10-15 cm in diameter and 5.0-7.5 cm high; this structure varies greatly among individuals and is visible
only when inflated and exposed by separation of the feathers. The iris is sepia-brown to greyish brown.
The bill is primarily horn-coloured to greenish yellow, mottled with black especially near the base. The
forehead, lores and bare skin of the head are mainly orange-red with numerous small black pigmented
spots, these increasing with age and most prevalent around the ear-opening and at the base of the bill.
The gular pouch and neck are pinkish or magenta, spotted with black especially on the pouch. The legs
and feet are nearly black but usually heavily coated with white excrement. There is a narrow band of
smooth reddish skin at the extreme upper end of the naked tibia, just below the beginning of the
feathers, visible only when the bird bends over or raises its wing.
When birds are together, the sexes are usually distinguishable by size. Males are larger than females,
with longer and heavier bills. Over 50 adults were measured in Uganda, and Pomeroy (1977b) found
that 'wherever the sum of wing-length and tail-length exceeded 100 cm, the bird was a male.
At the start of the breeding season upperparts become a paler bluish-grey, with a ladder-like band of
white marks along the upper surface of the wing, formed by the white edges of the greater secondary
coverts (Kahl 1987b). Luxuriant, elongated and fluffy undertail-coverts (the 'Marabou down' of the
former millinery trade) are pure white. The face and forehead become blackish red, encrusted with

what appears to be dried blood. Colours of the bare skin on
the head, neck and gular pouch intensify, with the back of
the neck becoming a bright pale blue. Iris colour, especially
in some females, turns a lighter brownish grey for several
days just prior to egg-laying (Kahl 1966a). The pale bluish
grey of the back is due to a 'waxy bloom' of keratin on the
surface of the feathers (Pomeroy 1977b, Jacob and Pomeroy
1979); this gradually wears oft'and the adults revert to the
darker slate-grey backs, usually long before the young have
fledged.
Hatchlings have wrinkled, pinkish skin and are sparsely
feathered with a pale grey down; newly hatched young
weigh about 70 g. Older nestlings of 8-35 days are covered
with a denser white down, except for the bare head, foreneck
and face; they are similar in appearance to nestlings of the
Greater and Lesser Adjutant Storks. Blackish contour
feathers appear on the back and wings at 35-50 days. At 51-
65 days, nestlings become predominantly black on the dor-
sal side, and after 66 days they are fully feathered and
change little in appearance thereafter (Kahl 1966b).
Juveniles are similar to adults. However, their uppersides
are dark slaty-black, with a moderate bluish gloss mixed
with brownish; they are also without the white hatchmarks
(secondary coverts) along the top of the wing. Denser pale
brownish or black filamentous feathers, mixed with pale
grey woolly feathers, grow on the head and neck. The soft-
parts are less brightly coloured, and the gular pouch is less
developed (or less commonly fully inflated). Bill-length in-
creases slowly with age, requiring 1-2 years to reach full
length (D. Pomeroy pers. comm. 1989). For a more com-
plete description of the development of plumages, and their
sequence, the reader is referred to Pomeroy (1977b).
The Marabou is generally a silent bird, except around the
nest and when roosting. During social displays birds clatter
Marabou Stork 135
their bills loudly and utter a variety of high-pitched squeals
and yelps, slurring to a lower-pitched, grunting cowlike
'Moo'; these vocalizations are quite similar to those given by
the Greater Adjutant Stork of Asia (Kahl 1966a, 1972a).
Their wings make loud 'Woofing5 sounds, especially during
landing and take-off during social encounters with other
Marabous.
Heavy on take-off, the Marabous flies strongly with noisy
rhythmic flaps once airborne. Groups soar on fixed wings in
thermals, sometimes climbing to such high altitudes that
they are nearly invisible from the ground. A group was
observed, from a light aircraft, soaring at 3000-4000 m
above Uganda with their gular pouches fully inflated (Akes-
ter et al. 1973). Colin Pennycuick (1972) provided an inter-
esting description of soaring behaviour in Marabous, as
observed from a motor-glider over the Serengeti Plains,
Tanzania. Unlike other members of the family, storks in
the genus Leptoptilos fly with their necks partially retracted
back onto their shoulders, similar to the herons, except
that the bill and legs are angled slightly below the horizon-
tal. We timed the flapping rate as 143/min (n = 8) (Kahl
1971b).
On the ground or at close quarters, the Marabou cannot
be mistaken for any other bird within its range. When
soaring on high, it can be distinguished from vultures and
the Great White Pelican (Pelecanus onocrotalus) by the long
legs trailing out behind, and from the rare Shoebill by the
thinner bill.
DISTRIBUTION AND POPULATION
The Marabou Stork is resident in tropical Africa. It occurs
from Senegal across to Ethiopia and Somalia, south to
northern South Africa (northeastern Transvaal, Zululand),
northern Botswana and northeastern Namibia.
The Marabou undertakes short, intratropical migrations
according to rainfall conditions. It often leaves areas during
the rains, to appear again the following dry season (Chapin
1932a:465, 467, Pomeroy 1973). In Uganda, migrant Mara-
bous seem to move mainly in a north-south direction
(Pomeroy 1977b). They also wander erratically to areas of
temporarily abundant prey. The species is a rare vagrant
south of its breeding range, and is an accidental visitor in
Israel (3 records; Paz 1987:34).
This species is common to abundant in most parts of its
range. Apparently the Marabou has increased in recent
years throughout its range, because of its close association
with humans. The total Uganda population was estimated
at 4000-5000 in 1970-72 (Pomeroy 1973) and had increased
significantly by 1975-76 (Pomeroy 1977c). The total
Kenyan population has been estimated at 1000-2000
(Pomeroy 1986).
The age-structure of the population in Uganda (based on
plumages of 4566 birds) is heavily dominated by adults:
13% juveniles, 15% subadults 3^ years old, and 71%
adults more than 4 years old. The maximum calculated age
exceeds 25years (Pomeroy 1977b). The estimated annual
mortality rate of adults is 8% per year (D. Pomeroy pers.
136 Marabou Stork
comm. 1990). The species has lived 31 years in captivity
(Flower 1938).
ECOLOGY
The Marabou is found mainly in open country, often in the
vicinity of large lakes or rivers. It does not occur in the
densely forested regions of Africa, except where forests are
bisected by large rivers. It seems to prefer arid or semi-arid
areas, but often within flying range of an aquatic habitat.
Often it is associated with vultures and other scavengers in
'big game country5, where it feeds on carrion provided by
the remains of kills left by large predators. It is common,
sometimes abundant, around human habitations, garbage
dumps, abattoirs and fish-camps, where refuse is available.
It also uses natural wetlands, especially during the nesting
season, and favours drying pools where fish are concen-
trated.
Feeding behaviour is surprisingly reserved. When feeding
at carrion with vultures and eagles, the Marabou is usually
a rather shy bird. It hangs around the outskirts of the group,
dancing in occasionally to steal a morsel from the smaller,
but more aggressive, scavengers. In such situations Mara-
bous play more the part of'pirate', allowing the vultures—
with their stronger, hooked bills—to tear off pieces of meat
from a carcass and then attempting to steal it away (Hous-
ton 1975).
At rubbish dumps and fishing villages, Marabous lose all
fear of humans and can be seen within a meter or so of
people working there, waiting patiently for a scrap to be
tossed their way.
Especially during the breeding season, but occasionally at
other times as well, Marabous forage for living prey. When
termites are swarming, the storks often stand at their
mounds and consume large quantities of these fat-bodied
insects as they emerge on their mating flights. Aquatic prey
is often hunted using Groping, the tactolocation technique
characteristic of the wood storks, Mycteria. The partially
open bill is immersed vertically in shallow water and moved
slowly about as the bird wades forward. When a prey item is
contacted, the bill snaps rapidly shut, and the prey is
quickly swallowed with a backward toss of the head. At
other times, when visibility is good, such prey is located
visually and Grabbed in the tip of the bill.
Marabous often feed together. In feeding groups of Mara-
bous, dominant individuals usually stalk about with their
gular pouches fully inflated, and subordinant birds give way
to them. Large numbers appear, as if from nowhere, at
infestations of locusts or army worm caterpillars and at
grass-fires, where they consume insects and small ver-
tebrates flushed or injured by the flames. Here they often
feed in the company of migrant White Storks and Abdim's
Storks. One Marabou collected while feeding on a swarm
of desert locusts (Schistocerca gregaria) had a total of 993
locusts in its alimentary tract (K.D. Smith and G.B. Popov
1953).
The food of the Marabou is almost anything organic.
According to Brown et al. (1982:188), it will eat Virtually
any animal matter from termites to dead elephant5. Over
200 Marabous have been recorded at one elephant carcass
in Kruger National Park, South Africa (Kemp 1974:15).
Carrion is an important staple in the diet, especially among
non-breeding Marabous. 'Wild5 carrion, such as the re-
mains of lion kills, has been largely replaced in many areas
by 'man-made5 carrion from garbage, fish scraps and abat-
toir refuse around settlements. And today in many urban
areas, vegetable matter forms a large part of the Marabou's
diet (D. Pomeroy pers. comm. 1990).
Whole vertebrates (primarily fish, frogs and rodents) are
favoured by breeding adults during the period that their
nestlings are growing most rapidly, probably owing to the
increased need for calcium by the young. Hand-reared nest-
lings did not survive when the supply of whole vertebrate
food failed (Kahl 1966b).
Marabous relish the eggs and young of crocodiles, which
they often consume as they are emerging from their shells
(Pitman 1957). Birds also sometimes fall prey to Marabous.
Wildlife film-makers Simon Trevor and Des Bartlett have
photographed them catching and eating nestlings of Quelea
quelea in Kenya's Tsavo National Park (see also Disney and
Marshall 1956, Leuthold and Leuthold 1972, and Kemp
1974) and adult doves in Namibia5s Etosha National Park.
Several authors tell of predation on larger birds, such as
young and adult flamingos in East Africa (Brown et al.
1982:188), cormorant nestlings in South Africa (Milstein
1975) and Great White Pelican chicks in Nigeria (Dunger
1965). With its massive bill and elastic throat, a Marabou is
able to swallow pieces of meat weighing as much as 600 g.
FJ. Jackson (1938:77) reported one 'struggling with the
entire windpipe of a bull buffalo that was halfway down its
throat5. A worker at the Kitale abattoir (Kenya), reported
that a Marabou stole and swallowed a bloody butcher-knife
that was laid aside for a few moments; days later the knife,
digested spotlessly clean of all blood and residue, was found
nearby in the grass, where it had been regurgitated as
'indigestible5 by the stork (Kahl 1988b).
It is estimated that an adult Marabou, weighing 5-8 kg,
requires about 0.7-1.Okg of food per day to survive (Kahl
1966b, Pomeroy 1973).
Often all the birds in an area gather in the late afternoon
at traditional communal roosts, which may be used for
many years if not disturbed. One such roost, near Nairobi,
Kenya, far from any known breeding colony, contained up
to 1000 individuals (D. Pomeroy pers. comm. 1990). The
next morning they often remain at the roost for several
hours after sunrise, until thermal air-currents develop on
which they can soar, effortlessly, to distant feeding grounds.
Marabous commonly sit for long periods on their tarsi and
often sun, either sitting or standing, with wings fully spread.
Opening the wings to the sun in this manner is a very
contagious behaviour pattern, and many birds in a group
may do so simultaneously.
The inflatable gular pouch is primarily used as a signal of
dominance in social displays. However, it and other sub-
cutaneous air-sacs may also serve other, secondary physio-
logical functions, such as thermoregulation.
Marabous are sometimes kept as pets in African villages,
and they become quite tame. However, their massive bill is

a formidable weapon, and they have been reported to have
killed children who were trying to catch them (Mackay
1950). In the Sudan, they are occasionally used as 'ani-
mated feeding perches' by the Carmine Bee-eater (Merops
nubicus) (Cunningham-Van Someren 1970).
BREEDING
Marabou Storks nest in colonies, often spread over a rather
wide area, of a few pairs to a several hundred pairs, most
commonly 20-60 pairs. They often build in association with
Pink-backed Pelicans (Pelecanus rufescens) or, less commonly,
with other waterbirds. Sites are frequently traditional and
are used for many years barring disturbance.
Consistent requirements for a successful breeding site
seem to be safety from disturbance, plus reasonable proxim-
ity (<50-60 km) to a good food-source (ideally both carrion
and live vertebrate prey) for the young. The largest colonies
in eastern Africa were near to extensive swamps (Pomeroy
1978a).
Nests are usually placed 3-40 m up in trees; they
occasionally nest on cliff-ledges, as is the case in north-
eastern Uganda, southern Sudan and southern Tanzania.
Tree-nests are often in euphorbia (Euphorbia dawei], mvule
(Chlorophora excelsa), baobab (Adansonia digitata) or acacia.
Such trees are sometimes in the midst of villages, especially
in West Africa (Bannerman 1930:107) or in larger cities, as
in Kampala, Uganda (Kasoma and Pomeroy 1987).
The nest platform is a rough structure of coarse sticks,
lined with smaller twigs and green leaves. Foundations from
previous years are often used again but usually require
considerable renovation, owing to weathering and stick-
pirating by neighbours. A finished platform is approxi-
mately 1 m in diameter and 20-30 cm thick. Occasional
sticks are added to the structure throughout the nesting
season, until the young have fledged.
Because nestling growth is so slow, the nesting cycle is
protracted: from egg-laying to fledging takes about 165 days
(Kahl 1966b). In the northern and southern tropics, breed-
ing usually begins in the dry season and young fledge during
the next wet season. In the equatorial zone, where there are
two wet/dry cycles per year, seasonality is less clear-cut
(Pomeroy 1978c); breeding cannot be completed in one
short dry season, and young may fledge in the next dry
season.
Pair-formation and ritualized displays in the Marabou
were described in detail in Kahl (1966a). Males establish
themselves on nest-sites and wait to be approached by
courting females. Their large gular pouches are usually fully
inflated at this time, indicating a high level of hostility and
dominance in the males. All trespassers are treated with
hostility at first; other males generally fight back, whereas
females react passively and with submissive displays, such
as the Balancing Posture. Eventually a persistent female is
accepted by the male.
After pairing the female also inflates her gular pouch, and
both members maintain them fully inflated much of the
time, at least up until the time of egg-laying. This seems
to signal a 'feeling of dominance', at the nest-site, in relation
Marabou Stork 137
to all other Marabous in the vicinity. When the pair is
firmly established, the male then periodically leaves her
alone on the nest-site while he flies out to collect sticks. A
mutual Up-Down is performed by the pair whenever one
member returns from an absence. Frequent copulations
follow, interspersed with nest-building and mutual display
activities.
During the early stages of nest-construction, the male
collects most of the material and delivers it to the female on
the nest, who does most of the building. Later in the breed-
ing cycle, a majority of the stick-collecting is done by the
female. Sticks continue to be brought to the nest by both
parents until the young are ready to fledge.
Clutch size ranges from 1-4, rarely 5, and usually 2-3.
The mean of 108 clutches was 2.4 eggs. Eggs are laid at
intervals of 1-3 days (Brown et al. 1982:190). The shells are
dull chalky white when first laid but are soon stained.
Both sexes share in incubation, feeding and guarding of
the young. Incubation begins at or soon after the laying of
the first egg. Both sexes incubate for about 84% of the
daylight hours: males 48% of the time, females 36% of the
time. Males also incubate more than females at night. The
incubation period is 29-31 days (average = 30.3 days). The
hatching process takes about 24 h. Eggs usually hatch at
intervals of 2-3 days; extremes observed are 1-7 days (Kahl
1966b, Pomeroy 1977b).
Parents take turns brooding the young much of the time
for first 10 days. Thereafter, brooding is reduced, except
during periods of rain or cold. Nestlings are shaded by a
standing parent, with wings either open or closed, when the
sun is hot. Young are almost never left alone on the nest
until after 25-30 days of age. Even then a parent is present
at or near the nest most of the time. Almost full-grown
nestlings are sometimes left alone while both their parents
forage. They are usually able to defend the nest against
intruding Marabous that sometimes attempt to enter it.
Nestlings solicit for food by dropping to their tarsi, with
partially open wings, cocked tail and up/down nodding of
head in the Begging display. Begging calls of newly hatched
young are a high-pitched chitter. This call gradually deep-
ens, as the young grow older, until it is a hoarse, nasal
braying.
Both parents feed the young, by regurgitation onto the
floor of the nest. We found that males fed more (55%) than
did females (45%) in 386 recorded feedings. Most feedings
occurred between lO.OOh and 15.00h, coinciding with the
daily thermal development that permitted the parent to fly
to and from distant feeding grounds.
We estimated food consumption of young, from captives.
It increases from <100g/day at age 1-10 days, to 500-
600 g/day at age 60+ days. Nestlings receive a large pro-
portion of whole-vertebrate food, such as fish, probably
because of their high calcium requirements during rapid
growth. Of 421 feedings observed at the Kitale colony
(Kenya), 29% consisted of whole-vertebrate prey (Kahl
1966b).
Larger nestlings are also given water, by regurgitation, by
their parents. This watering may be related to the thermo-
regulatory habit of excreting a dilute urine onto the bare
138 Marabou Stork
skin of their legs for evaporative cooling (Kahl 1963b).
Without a supplementary water supply, from the parents,
this behaviour might lead to dehydration of the young.
Nestlings can stand and flap their wings at 17 days old,
hover over nest at 65 days, and make short flights within the
nest-tree at 95 days. The first recorded flight out of the
tree was at 98 days, although more commonly first flights
were at 110-115 days. Nestling growth is very slow, and
young remain dependent on their parents until an age of 130
days or more. Such a protracted growth pattern is perhaps
an adaptation to reduce the nestlings' daily food demands
to a level the parents can meet (Kahl 1966b, Pomeroy
1977b).
Marabous are probably 6 or 7 years old when they first
breed (Pomeroy 1977b). Some adult Marabous may not
attempt to breed in any given year and among those that do
breed, success is fairly low. Depending on conditions, 52-
95% of eggs hatch. In nests where 3 eggs hatch, it is rare for
the last-hatched chick to survive, although it occasionally
does (Pomeroy 1979). Only 36% of eggs in the Kitale colony
(Kenya) resulted in fledged young (Kahl 1966b). About 0.9
young per nest fledged in Uganda colonies (Brown et al.
1982:190). In recent years (1984-89), the Marabou has
increased within the city of Kampala, Uganda, and breed-
ing success has risen to 1.18-1.75 young/nest (D. Pomeroy
pers. comm. 1990).
Pomeroy (1978a) found that nesting success was greatest
in the medium sized colonies (i.e. 40-60 nests). The reasons
for this are unknown. It may be that medium-sized colonies
had a larger proportion of mature, experienced adults and,
therefore, had a higher success rate. According to the calcu-
lations of Pomeroy (1978b), only about one egg in three
[gives] rise to fledglings .. . less than a quarter of these
survive to join the breeding population. Hence any extra
recruit to the breeding population would require 12 extra
eggs [to be laid].
TAXONOMY
The relationships of the Marabou to the Greater Adjutant
Stork have been a matter of some discussion, owing to their
similarity. However, we have found marked differences in
their behaviour, sufficient to continue to recognize them as
separate species. We discuss this further in our account of
the Greater Adjutant Stork.
CONSERVATION
At the present time, the Marabou Stork appears to be doing
very well indeed in most parts of its range. It is one of those
unique species that appears to benefit greatly from its
association with man and his urban environment. It is
interesting and puzzling that this species is increasing in
Africa, when its close relative, the Greater Adjutant Stork, is
decreasing in Asia.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 304.
 Shoebill
Balaeniceps rex Gould

Balaeniceps rex Gould, 1850, Athenaeum, no. 1207, p. 1315: Upper White Nile (=Sudan)

Other names:
Whaleheaded Stork, Boat-bill, Bog-bird [Sudan], Lesser lechwe-eater [Zambia] (English);
Schuhschnabel, Konig Walkopf [='King Whalehead'] (German); Bec-en-sabot du Nil (French);
Becco a scarpa [Italian]; Schoenbekooievaar (Dutch); Abu markub [='Father of the shoe'] (Arabic);
Adungwek or Adumwueg (Dinka); Bany (Nuer); Okomo warjok (Shilluk); Bulue (Luganda);
Mkumabimo or Nkubabimu (Bakenya/L. Kyoga, Ugandan); Ekurududu (Teso/Ugandan);
Chifumpafumpa (Tabwa/Zambia); Fumpa-fumpa (Balamba/Katanga, Zaire); Motuta or Mututa
(Lualaba/Katanga, Zaire)

IDENTIFICATION
Slow moving, deliberate, morose, often statue-like, the Shoebill is a large grey bird, standing 110-
140 cm tall. It is immediately recognizable by its huge bulbous bill, reminiscent of a Dutch wooden clog.
The adult's head is dark grey, with a slightly darker blue-grey crown, while its back and mantle are
ashy grey with a slight greenish gloss. The lower neck feathers are lanceolate, with black shafts. The
primary wing feathers are dark grey, although black near the tips. The secondaries have a slight
greenish gloss. The underparts are paler grey. The enormous, prominently hooked bill is yellowish with
dark irregular patches. It is a formidable tool and weapon, with cutting edges like a turtle's jaws (FJ.
Jackson 1938:65). A dark keel on the upper mandible terminates in the hook, which covers the tip of the
lower mandible when the bill is closed. The eye is a pallid yellowish or greyish white, and the legs
blackish to horn-coloured. The toes are extremely long (middle toe = 16.8-18.5 cm long; W. Fischer
1970:13) and completely divided (i.e. not partially webbed, as in the true storks); the claw of the hind
toe is larger than those on the fore-toes (Reichenow 1900-01:357). A small crest or tuft on the back of
the head can be erected during alarm or excitement (Owen 1958, M. Burton and C.W. Benson 1961,
C.W.Benson 1961).
The male is noticeably larger than the female and has a longer bill (see Abb. 14 and 15 in W. Fischer
1970:20, and Figs. 4 and 5 in L. Buxton et al. 1978).
The breeding plumage is not known to differ noticeably from the regular adult plumage. The eye may
become more yellow at sexual maturity.
Newly-hatched young are covered with a silvery grey, silky down. The bill, though not yet bulbous, is
markedly hooked and the gape is wide; the head is large relative to the body. Exaggerated swelling of
the bill becomes noticeable at 23 days and is well-developed by 43 days (L. Buxton et al. 1978). Later,
'nestlings are covered with grey down of a fulvous tinge' (A.L. Butler 1905). The iris colour of
immatures is said to change from milky blue-grey, to pearl-grey (in the second year), to bluish white (at
about 5 years), to pale yellow (perhaps at sexual maturity) (W. Fischer 1970:10).
A chronology of the development of the nestling is given by L. Buxton et al. (1978): Days 1-35,
entirely or mainly downy, unable to stand, requires constant brooding or shading; days 35-75,
developing contour feathers, stands unsteadily or sits on tarsi, requires little brooding by day; days 75-
95, largely feathered, left alone by parents, seldom shaded, stands steadily and exercises wings, begins
to leave nest on foot, soon can fly.
When fully feathered, juveniles are a darker grey than the adults with a slight brownish tinge. The
bill develops its bulbous shape but is not quite as exaggerated as the adult birds. In one Zambian

juvenile, it was a pinkish colour with ill-defined blackish
markings (L. Buxton et al. 1978). Plumage parts that are
slate-grey in adults are grey-brown in juveniles (Reichenow
1900-01:357, Schouteden 1912). Captives assume adult plu-
mage at 3^ years (Moller in Collar and Stuart 1985:66).
During displays at the nest, the bill is clattered and a high
pitched whine or 'Moo' is uttered (W. Fischer 1970:45, 58,
L. Buxton et al. 1978). A soaring bird uttered a 'rather loud,
hoarse croak' as it dived at a soaring Marabou Stork nearby
(pers. obs. M.P.K.). Also reported are a hoarse but feeble
cry (Dekeyser 1962), a raucous and guttural cry
(Dechambre 1936) and an 'occasional loud cry while soar-
ing' (Mackworth-Praed and Grant 1970:51). Both adults
and young clatter their bills (Gilliard 1958). Recordings of
the voice are available (Boswall and North 1967). Begging
calls of nestlings resemble human hiccups (L. Buxton et al.
1978).
Near-vertical take-offs and landings are possible, because
of the large, broad wings (Guillet 1979). When disturbed,
this stork is able to rise more calmly than any of the larger
herons, without noise, and flap slowly away (FJ. Jackson
1938:65). The neck is retracted in flight (Schouteden 1912,
Bouet 1955a). It does not readily take flight and is unlikely
to travel more than 400-500 m when flushed (Owen 1958).
It soars on thermals (Berg 1940). On 8 December 1963, a
single Shoebill was watched soaring high up with several
Marabou Storks just soutwest of Kampala, Uganda. The
wings were held perfectly flat (i.e. no dihedral) and the
primaries were widely slotted; the extremely long toes were
conspicuous in flight. The flap rate has been timed once; at
150 flaps/min (pers obs. M.P.K.) it is slower than all but the
largest of storks (Kahl 1971b).
The Shoebill is unmistakeable if seen well. In flight or
when soaring, it might be mistaken for a Marabou Stork or
possibly for a Goliath Heron, but the Shoebill has a shorter,
blunt bill, whereas the heron's and Marabou's bills look
Shoebill 141
sharp and needle-like, especially from below, (pers. obs.
M.P.K.).
DISTRIBUTION AND POPULATION
The Shoebill is widely but locally distributed in freshwater
swamps of central tropical Africa, from southern Sudan
(north to about 9°N) through parts of eastern Zaire,
Rwanda, Uganda and western Tanzania to northeastern
Zambia. The distribution of this species generally coincides
with that of papyrus (Dekeyser 1962), and it also largely
coincides with that of the lungfish (Protopterus aethiopicus)
(Greenwood 1958:14).
The Shoebill is most numerous in the White Nile Sudd
and adjacent areas of the southern Sudan; significant popu-
lations also occur in the wetlands of Uganda and western
Tanzania and the Bangweulu Swamp of northeastern Zam-
bia. In Uganda, it is widespread—but with a fragmented
distribution in suitable habitat—eastward to Teso and
Busoga, and south to the Kagera river. Records of its occur-
rence in the extreme west of Kenya (A. Koenig 1910, Brit-
ton 1978, 1980) are apparently erroneous (I.S.C. Parker
1984a). It has been found in Tanzania at several sites, all in
the west [Moyowosi Swamp, Lake Mujungwizi, Lake
Mujunju, Kagera Swamp, Nzilindagaza, Malagarasi
Swamp, and Lake Rukwa (rarely)]. In Zambia, we have
isolated records from the Itawa Swamp, near Ndola, and
the Kasanka National Park; but in the west of the Northern
Province and in the Luapula Province, it is more wide-
spread, in particular in the Bangweulu Swamps. It is also
reported uncommonly from Fungwa Swamp, around
Mweru Marsh, Lake Kako, and near the mouth of the
Luapula River. In Rwanda, it occurs in the Akagera de-
pression. In Zaire, it is found in the western Rift Valley
(Lake Edward) and mainly in the southeast, in Katanga
Province.
Isolated birds are reported or rumoured from the Central
African Republic (Blancou 1939), northern Cameroon
(Lisle 1956), southwestern Ethiopia (Duckworth 1974),
Malawi (K. Brouwer pers. comm. 1989); and strays have
been reported as far southwest as the Okavango Delta,
Botswana (N.J.C. Mathews 1979, pers. comm. 1989). Long
ago it was rumoured on the headwaters of the Upper Congo
River, in Angola (H. Johnston 1902). These and other
distribution records are discussed in Collar and Stuart
(1985:59-63).
Drawings of the Shoebill dating from approximately 3500
BC, have been found in Egyptian tombs (A. Koenig
1910:525; Schouteden 1912). Fossil relatives of the Shoebill
have been found from the Oligocene of northern Egypt
(D.T. Rasmussen et al. 1987) and the late Miocene of
Tunisia (Rich 1972).
The Shoebill appears to be non-migratory, remaining in
or near its home range throughout the year (Kasoma and
Pomeroy 1987), so long as foraging conditions are appropri-
ate. Sudanese birds make limited seasonal movements, in
response to changes in food availability, habitat availability
and disturbance (Guillet 1978b, and pers. comm. 1990).
It is generally thought that this is a rare species, its
142 Shoebill
population being limited mainly by the distribution of its
habitat. It may be that isolated pockets of birds or of
individual pairs exist in suitable areas throughout its range.
Even in areas where it is relatively numerous, estimates of
numbers can vary considerably. As I.S.C. Parker (1984b)
stated,
an impression of scarcity probably derives from the
general inaccessibility of the species' habitat. In the few
areas where it can be searched from a boat or from a
vehicle, the habitat is usually a thin ribbon along the
water's edge. In such circumstances the bird's scarcity
is not surprising. Where the habitat is extensive, in the
shallows between dry land and papyrus banks, it is
difficult to search from either boats or wheeled vehicles.
In order to find it 'its home must be entered; cruising to and
fro outside ... would never reveal... that it may be ... just
beyond the barrier of papyrus'(F.J.Jackson 1938:66).
In the Jonglei area of the southern Sudan, 3000 were
counted in 1979 along the White Nile system (A. Guillet in
Collar and Stuart 1985:63), and Jonathan Kingdon (in
Collar and Stuart 1985:63) estimates that the whole area of
the Sudd may contain as many as 10000 birds. In Rwanda,
numbers have fallen from 30-50 in the early 1970s to be-
tween 15-25 (Vande Weghe 1981), and in Uganda the
population has been estimated 400-600 birds (Collar and
Stuart 1985:64). However, Kasoma and Pomeroy (1987)
suggest that this estimate may be too low, since the birds are
difficult to see, and Moller (1978) found 26 birds in 20 km2
of swamp in Uganda. I.S.C. Parker (1984b) estimated
about 0.64 birds/km2 in a swamp in western Tanzania,
surveyed from the air. In a portion of the Bangweulu
Swamps of northeastern Zambia, G.W. Howard and D.R.
Aspinwall (1984) estimated that 232 (c. 0.145 birds/km2)
Shoebills were present in October 1983. A reasonable esti-
mate of the total number of Shoebills would seem to be
between 11000-15000 birds over its whole range (Anon.
1985b, A. Guillet pers. comm. 1989); this is nearly 10 times
greater than previous estimates (Guillet 1978a, Brown et al.
1982:191).
ECOLOGY
Shoebills are characteristically birds of extensive, dense
freshwater marshes, especially flood-plain interspersed with
undisturbed papyrus (Cyperuspapyrus) and reedbeds (Phrag-
mites, Typha, etc.) having floating vegetation on which to
stand; they generally avoid areas in which the plants are
taller than the bird's back or are too dense to penetrate
easily. They seem to prefer areas of poorly oxygenated
water, where fish and other aquatic organisms are forced to
the surface to breathe (Guillet 1987:222). Because of this,
Vande Weghe (1981) found that, in the Akagera National
Park and probably elsewhere, the Shoebill avoids pure
papyrus swamps, preferring mixed swamps of Cyperus papyr-
us (papyrus), Miscanthidium and floating vegetation consist-
ing of ferns and grasses with clumps of Cladium and Typha. It
is also found in marshes dominated by Pistia stratiotes, Typha
(cattail), Miscanthidium., Vossia cuspidata, Cyperus latifolius and
Echinochloa pyramidalis. When conditions are favourable,
birds have been seen on flooded grassland and open flood-
plain (Collar and Stuart 1985:64).
In the southern Sudan, the Shoebill has recently
expanded its feeding habitat to include rice fields and other
flooded plantations (A. Guillet pers. comm. 1990). This
species especially prefers open parts of the swamp (Owen
1958), possibly due to the ease of taking flight under such
conditions, and also because in dense papyrus swamps fish
are harder to see and capture.
Shoebills feed in shallow or deep water by Standing and
waiting or Walking Slowly, usually on floating vegetation or
along channels (L. Buxton et al. 1978, Guillet 1979). These
feeding methods are the 'lurking and stalking' (Lauern und
Pirschen) described by Moller (1982a) and by Guillet (1979).
'Instead of searching for his prey, [the Shoebill] waits
patiently for it to come to him' (A.L. Butler 1905). Ambush
is their speciality.
Standing in one place, motionless, may last for 20-30 min
(L. Buxton et al. 1978, Brown et al. 1982:192), but at other
times Standing and Walking tend to blend together. Walk-
ing tends to be used more commonly in sites where uninter-
rupted movement is possible, such as along channels. On
floating platforms of vegetation, feeding is restricted to
Standing and waiting (Guillet 1979); it uses its extremely
long toes to support its weight on floating plants—much in
the manner of a giant jacana—often sinking slowly up to the
tarsal joint or even deeper. It does not habitually wade in
deep water, nor stalk prey in a crouched heronlike posture.
Hopping is observed only as a startle reaction and it does
not often run, although we once saw a bird run a few steps
after something, with wildly flapping wings (pers. obs.
M.P.K.). Normal progress is with a very slow stride, each
step taking on average 5 s, and carrying the bird forward
about 60cm (Moller 1982a). Occasionally it may briefly
adopt a crouched posture during pursuit of a prey item.
Owing to these preferred feeding behaviours, channels,
lagoons and waterways that carry comparatively slow-
moving water are used for feeding. Especially favoured
feeding places are along trails made by large animals, which
open up areas that would otherwise be too thick for Shoe-
bills to utilize.
Fishing is by visual location, rather than by touch, and
hearing may also play a role (Guillet 1979). The bird feeds
by holding the bill pointed vertically downward, to facilitate
binocular vision. Fish, such as lungfish, are captured as they
come to the surface to breathe, particularly in stagnant (i.e.
deoxygenated) water. The final attack is performed ex-
tremely rapidly and, apparently uniquely, by 'collapsing' on
the prey, throwing the wings forward and plunging down
with the head and bill (Guillet 1979, Moller 1980, 1982a).
The bird sometimes becomes entangled in vegetation, tak-
ing some time to recover. The tip of the bill absorbs the
major force of the bird's relatively violent strike (Guillet et
al. 1985), and the prey is secured by the sharp hook on the
upper mandible. Vegetation, which is often taken into the
bill along with the prey, is separated (Guillet 1979), and the
prey is often decapitated by the extremely sharp mandibles
before swallowing. From 1 to 10 min may elapse between

capture and swallowing of prey; turtles and snakes may take
even longer (Moller 1982a). After the prey has been swal-
lowed, water is often scooped up in the bill for drinking. On
rare occasions 'collapsing' may be done from on the wing
(Guillet 1979).
The Shoebill's unique, boat-like bill is highly adapted to
the bird's specialized mode of feeding (Guillet et al. 1985,
Guillet 1987). Thanks to the anatomy of the bill and related
skull structures, the Shoebill is able to catch unwieldy prey
amid dense aquatic vegetation. A unique anatomical lock-
ing mechanism in the articulation of the jaw (Bock 1964:18)
probably assists the bird in holding onto heavy, hard, or
slippery prey. There is no evidence to confirm the rumour
that Shoebills excavate aestivating lungfish from dried mud
(Brown et al. 1982:192).
Shoebills tend to forage singly, even if otherwise paired
(Moller 1982a), seldom closer than 20 m to each other. We
once saw 9 birds, scattered over a large marsh, southeast of
Lira, Uganda (pers. obs. M.P.K.). Loose aggregations
occur, particularly where water is drying out and large
numbers offish have become concentrated (Guillet 1979).
There is no evidence that Shoebills ever fish communally, in
the manner of some pelicans, as suggested by P. A. Cottam
(1957). Moller (1982a) reported that, in Uganda, most
feeding takes place during peak periods between 06.00 and
08.00 h in the morning, and between 18.00 and 20.00 h in
the evening. Reports of night feeding are often discounted
(Brown et al. 1982:191), although Guillet (1979) reported
that they can feed by the light of fires set by fishermen, and
their very large eyes would suggest some noctural, or at least
crepuscular, abilities (FJ. Jackson 1938:66). W. Fischer
(1970:50) refers to the Shoebill as a Dammerungsvogel
( = twilight-bird). A bird in the marshes bordering Lake
Edward, in western Uganda, was followed on all of 3
consecutive days (Moller 1982a); it moved approximately
0.5-1.5 km during foraging activities each day but appar-
ently did not move from its resting place at night. On the
other hand, C.R.S. Pitman (in M. Burton and C.W. Benson
1961), who had considerable experience with the species,
reported that they feed 'mostly by night' and that they
become active in the evening.
In 551 h of observation in Uganda, Moller (1982a)
recorded 111 strikes, 60 of which (54%) were successful.
The shortest time between successful strikes was 23 min,
and birds usually changed locations after a strike (Moller
1982a). This may indicate that the disturbance of a strike
makes another successful one unlikely at the same place
soon after. It seems that birds are fortunate to make about 2
kills per day.
The diet of Shoebills is primarily composed of fish, espe-
cially lungfish (Protopterus aethiopicus), bichirs (Polypterus sene-
galus), catfish (Clarias spp.) and tilapia (Tilapia spp.). They
also eat frogs, watersnakes, monitor lizards (Varanus niloti-
cus), turtles and young crocodiles (Crocodilus niloticus).
Rodents, snails and young waterfowl are taken on occasion.
It has been reported that Shoebills will take newborn lechwe
(Kobus leche) calves, but such reports (M. Burton and C.W.
Benson 1961) are probably without foundation. (L. Buxton
et al. 1978, Brown et al. 1982:192), especially since lechwe
Shoebill 143
calves are large and Shoebills swallow their prey whole (A.
Guillet pers. comm. 1990).
In any case, fish provide the bulk of the diet, and in many
areas these include impressively large lungfish. In Uganda,
Moller (1982a) observed that fish 15-50 cm long were eaten;
a lungfish 1 m long was caught but then released. In areas
frequented by hippopotamuses, the cloudy water may
enhance the accessibility of large fish (Moller 1980). Ideal
feeding sites have high densities of fish but are low in
dissolved oxygen, causing the fish to rise for air.
The diet does vary geographically. Moller (1982a)
reported that lungfish and catfish are the main prey taken in
Uganda, whilst L. Buxton et al. (1978) found that catfish
(Clarias mossambicus) and water snakes were the main diet
fed to young in the Bangweulu Swamps, Zambia. Zambian
birds caught catfish weighing up to about 500 g and snakes
50-60 cm long (L. Buxton et al. 1978:205). A captive juven-
ile consumed approximately 0.91 kg of fish and meat per
day (G. Steinbacher 1937, Owen 1958).
The Shoebill's usual stance is with the bill resting on the
breast (Mackworth-Praed and Grant 1970:51). It habitually
stands motionless and statuelike for many minutes at a time,
apparently waiting patiently for a lungfish or other prey to
surface; it was characterized by FJ. Jackson (1938:65) as
'sombre-looking and listless .. . often looking more like a
grey stump or post than a bird'. Often it stands on one leg
while scratching with the other (Reichenow 1900-01:358,
Guillet pers. comm. 1990). It does perch in trees, although
none exist in much of its habitat (A.L. Butler 1905, M.
Burton and G.W. Benson 1961, Guillet 1984).
Shoebills soar on hot afternoons. At times, at least, this
may have an advertising and territorial function (L. Buxton
etal. 1978, Moller 1980).
At least in some areas neither nesting nor fishing birds
can tolerate even a low level of human disturbance (Guillet
1978a). They will desert a breeding area if disturbed before
egg-laying (A. Guillet, pers. comm.). When the surrounding
vegetation had not been burned and heavy cover remained
intact, birds at established nests tolerated native fisherman
as close as 20 m (Guillet 1984). Other observers have also
commented on the lameness of the birds after the nest has
been established (L. Buxton et al. 1978).
Captive birds in the Tierpark Berlin periodically shed
some of the covering on their bill (W. Fischer 1970:80-81);
it is not known whether birds in the wild also do this. Also in
the Berlin Tierpark, we have seen a captive Shoebill sun-
bathe, with its back to the sun and its wings spread with the
tips touching the ground; when a cool wind started, the bird
closed its wings (pers. obs. M.P.K.).
BREEDING
In Sudan, Shoebills nest solitarily, usually with less than
3 nests/km2. An area around the nest is often cleared by the
adult birds of all tall vegetation for an area of about 3 m in
diameter (Guillet 1984). In Uganda, a breeding pair may
defend a 2-4 km2 area from intruders (Moller 1978).
In the northern and southern extremes of its range, nest-
ing begins after the rains end, just as water levels begin to
144 Shoebill
drop. At both Bangweulu, Zambia, and the White Nile
region, Sudan, eggs are laid at the end of the rains, and
young fledge late in the dry season or early in the next rains
(Brown and Britton 1980:38). The very long breeding cycle
of the Shoebill forces it to lay eggs in a wet season in some
equatorial areas that have two rainy seasons per year. This
is the case in Uganda, where Shoebills nest in March-June
during the main rains (FJ. Jackson 1938, Kasoma and
Pomeroy 1987). Breeding at this time would produce young
fledging in the main dry season.
Both birds build the nest, performing Up-Down greeting
displays often in association with collection and presen-
tation of nesting material (L. Buxton et al. 1978). Up-Down
displays accompanied by bill-clattering are the commonest
displays seen at the nest (Guillet 1984). Both birds stand
erect with bills pressed close against the neck. They bow,
point the bill downward and shake it repeatedly from side to
side, accompanied by bill-clattering. The bill-clattering is
rather slow (4-5/s) and has a deep, hollow quality, clatter-
ing sounds of male and female are individually distinguish-
able (Brown et al. 1982:191), probably owing to the relative
sizes of their bills. Sometimes, at the beginning of an Up-
Down, the bird gives a hoarse, rasping squeal, then the bill
is lifted to about 45° above the horizontal, so that the back of
the head and crest rest on the upper back; then it brings the
head down while clattering the bill (W. Fischer 1970:45, 58,
pers. obs. M.P.K.). Vocalizations are also described as a
high pitched whine or 'Moo' (L. Buxton et al. 1978). The
Up-Down shows some apparent similarities to the display
of the Marabou Stork (Kahl 1965, 1966a). Up-Downs are
also given as a greeting to familiar people by captive birds
(A.L. Butler 1905, G. Steinbacher 1937, Poulsen 1950, W.
Fischer 1970).
Other displays described by W. Fischer (1970) also re-
semble stork displays more than they do those of herons or
pelicans (other suggested relatives of the Shoebill —see Tax-
onomy).
In most areas pairs seem to defend all-purpose territories,
2.5-3.8 km 2 in size, with the nest fairly centrally located
(Moller in Collar and Stuart 1985:66). Adults on the nest
defend it with displays similar to the Forward Display of the
storks, and also with bill-clattering and a display termed
Bill Hook Exposure, in which the bill is raised, opened and
pointed at the intruder. If ritualized threats do not intimi-
date an intruder, a bird 'jumps into the air and falls feet first
onto its rival; the bill then follows, with the whole body
weight behind it' (Guillet 1984:234).
Nest-building is by both sexes and, before the eggs are
laid, involves the birds 'sewing' new grass into the structure
by jumping up and down and working the material in with
their toes. After the eggs are laid, new additions of grass are
just dropped on the nest, without any 'sewing' movements
(Guillet 1984). The description of the behaviour of the
Shoebill when collecting or adjusting nest material sounds
somewhat similar to the Swaying Twig-Grasping display in
Marabou Storks (Kahl 1966a, L. Buxton et al. 1978:218).
The nest is usually built on a floating platform, in exten-
sive marshes, and surrounded by high emergent vegetation
and sometimes deep water (Guillet 1978a), sometimes it is
in an open, grassy glade within stands of dense papyrus
(C.W. Benson 1961). Nest-mounds are large and flattish, up
to 3 m across, built of floating vegetation, which may be
partially submerged. The actual nest, often constructed of
grass stems and measuring c. 1.0—1.7 m across, is placed on
top of this mound. In Sudan, floating Shoebill nests will
often support the weight of a man (Guillet 1984), but such
was not the case at a floating nest in Zambia (L. Buxton et
al. 1978). The report (Brown et al. 1982:192) that nests
become water-logged and sink during the nesting season is
incorrect (A. Guillet pers. comm. 1990). In some areas of
the Sudan, nests are placed on termite mounds projecting
about 1.5m above water level (C.W. Benson 1961, Critch-
ley and Grimsdell 1970, Guillet 1978a). Nests are often
added to and reused in successive years (Guillet 1984).
The white eggs have a thick chalky coating and become
much stained and dull during incubation. The normal
clutch size is 2 eggs; 1- and 3-egg clutches do also occur
(average = 1.7, n — 7; L. Buxton et al. 1978). Chicks can
hatch with as many as 5 days between them, so that the first
egg probably is incubated immediately upon being laid.
Eggs (and later young) are regularly wetted with regurgi-
tated water on hot days, up to four times daily. Eggs are
turned either by the bill tip or by pressing the grass around
the eggs with the toes. L. Buxton et al. (1978) considered
this behaviour to be intentional rather than accidental; we
have not seen egg-turning with the toes in any of the true
storks. Both parents incubate. Nest-reliefs at a Zambian
nest usually came in mid-morning and again in the after-
noon, with the female doing about 75% of the incubation,
including the night hours (L. Buxton et al. 1978). The
duration of incubation is not known, but it is probably
about 30 days.
Nestlings are brooded, shaded, guarded and fed by both
parents; however, at a Zambian nest the female was the
more attentive (L. Buxton et al. 1978). The chicks give a
hiccuping Begging Call, and peck at the legs of the adult
and the hook on their bill-tip, thereby stimulating regurgi-
tation. Adults hold the food, which is well champed up into
soft pulp, between the mandibles from which the chick can
obtain it; this mode of feeding is continued until whole fish
can be swallowed by the chick, unaided, at about 30 days.
Any food dropped onto the floor of the nest is later re-eaten
by the adult.
This method of feeding young differs from true storks,
who regurgitate directly onto the floor of the nest. During
the last "6 weeks of its time in the nest, a single Zambian
young was fed an average of 6 fish and/or snakes per day (L.
Buxton etal. 1978).
Small chicks are continually shaded by the attending
parent and on hot days regularly doused with water; some of
the regurgitated water is drunk by the chicks. Shading
continues until feathers are fully developed at about 60 days
(L. Buxton et al. 1978). The pre-fledging period lasts about
95 days, at which time young first leave the nest on foot.
They can fly at about 105 days and fly well by 112 days, but
are still fed by the parents for a further period—possibly a
month or more. Thus, from egg-laying to first flight takes
135-140 days, according to Guillet (1978a:42) and L. Bux-

ton et al. (1978); however, the latter authors stated that 'in
all, from the start of nest-building to independence of the
young, the nesting cycle would occupy 6 to 6i months' (180-
195 days). Most nests have been reported to contain 1 or 2
young, occasionally as many as 3 (A.L. Butler 1905, L.
Buxtonetal. 1978).
TAXONOMY
As discussed in the chapter on classification, there has been
historical debate on the affinities of the Shoebill. Powder-
down patches on the lower back suggest possible affinities
with the herons (Bartlett 1861), tongue morphology is simi-
lar to the Hamerkop (Scopus umbretta), and the syrinx is
heron-like (Beddard 1888). The structure of the skull has
led some authors to conclude that the Shoebill is a heron
(Boehm 1930) .and others to suggest a relationship to the
pelicans (P.A. Cottam 1957, Saiff 1978). Even in the last
century, von Heuglin (1869-73:1096) called it 'a Marabou
. . . with a pelican-like bill'. However, we feel that the
structure of the bill—and related features of the skull—are
so closely tied to the bird's specialized foraging techniques
that they are probably not a reliable indication of ancestral
relationships.
Some recent studies, both behavioural and morphologi-
cal, have converged on the conclusion that the species is
more closely related to storks than to any other group (A.
Feduccia 1977, L. Buxton et al. 1978). However, based on
recent biochemical analyses, Sibley et al. (1988) have classi-
fied it as a subfamily of the Pelecanidae (with the true storks
sharing an adjacent family with the New World vultures).
The question as to whether the Shoebill excretes a dilute
urine onto its legs in reaction to hyperthermia, as do all true
storks (Kahl 1963b), is still unsettled. Heat-stress trials
(Kahl 1967a) resulted in no urohydrosis by a captive Shoe-
bill. However, L. Buxton et al. (1978) present some evidence
[not sure proof, as suggested in Brown et al. (1982:190)] that
Shoebills may at times excrete on their legs as a cooling
mechanism. Watering of nestlings by parents may be related
to this behaviour, since other similar birds (e.g. herons) are
not known to do it. Because urohydrosis is probably rather
complicated, from a physiological and genetic standpoint, it
may be valuable in assessing taxonomic relationships.
Clearly more work needs to be done on this problem.
The Shoebill is distinctive in many aspects of its biology,
and is thus, we feel, most appropriately placed in its own
family, the Balaenicipitidae (L. Buxton et al. 1978, Kahl
1979a). The fact that we have included it in this book, while
excluding the Hamerkop, indicates that we believe the
Shoebill to be closer to the storks (Kahl 1967b, 1979c);
however, the jury remains out as to how close.
CONSERVATION
The highly specialized ecology, patchy distribution, small
populations sizes and extreme dependence on marshland
that is subject to disturbance and drainage renders the
Shoebill a highly sensitive species. The ICBP rates its posi-
tion as of'special concern' (Collar and Stuart 1985:59). It
has been included in Appendix II of the CITES Convention
(K. Brouwerpers. comm. 1990).
Shoebill 145
The main area of concern for this species is undoubtedly
the southern Sudan. Major dangers are large-scale drainage
(especially for the Jonglei Canal, irrigation and rice farm-
ing), cattle grazing and burning of trails, and increased
fishing activity. All of these factors threaten Shoebills, espe-
cially when water levels drop late in the breeding season and
nests are particularly vulnerable to cattle, fire and humans.
In some areas, boundaries between Sudanese tribal hold-
ings create a sort of 'no-man's land', in which Shoebills can
sometimes nest undisturbed (Guillet 1978a). A major con-
cern in Sudan was the planned construction of the Jonglei
Canal; however, civil war has halted work on this project, at
least temporarily.
There is a demand for the Shoebill by zoos, because of its
large size and distinctiveness. Brown et al. (1982:193) stated
that 500 young were reportedly taken in 1978, but all of
these birds were said to have perished in the Khartoum Zoo
(K. Brouwer pers. comm. 1990). By late 1990, Shoebills
were being offered for sale in the USA for US$10 000-20 000
each, making them the most expensive of all the available
birds in the zoo trade (Bill Toone, San Diego Wild Animal
Park, pers. comm. 1990). Captive breeding has apparently
been reported only once, in 1964 in Elizabethville
( = Lubumbashi), Zaire (Jarvis 1966); since then, about 100
birds have been kept in zoos in the past 20 years, with no
reproduction known (Collar and Stuart 1985:70, Guillet
1987, R.E.Johnson et al. 1987a).
Not only does the taking of birds for sale reduce wild
populations, but it teaches local people that such birds are
extremely profitable, and encourages hunting and conse-
quent disturbance. Many tribal customs forbad the hunting
of Shoebills, which were feared and protected by myths and
legends (M. Burton and C.W. Benson 1961). For example,
people around Lake Salisbury, Uganda, believe that when
in a canoe, you must never mention the name of the Shoe-
bill, otherwise there will be a storm and the canoe will
overturn (L. Tennant pers. comm. 1964). The policy of
buying Shoebills from local people is helping to destroy
these protective beliefs. Other people, not bound by such
customs, are also penetrating these areas (Guillet 1978a).
We feel that only zoos with serious breeding programmes for
the species should be encouraged to purchase Shoebills.
The Shoebill is protected by law in Sudan, the Central
African Republic, Uganda, Rwanda, Zaire and Zambia
(Collar and Stuart 1985:67). It is included in Class A of the
African Convention of Nature and Natural Resources (K.
Brouwer pers. comm. 1990). However, the construction of
large water projects will potentially upset the ecological
balance in several crucially important areas. Should the
three-phase project envisaged for the eventual re-routing of
the White Nile (the Jonglei Canal) be undertaken, the
flooding pattern in the Sudd region of the Sudan would be
completely altered, destroying the habitat. In Uganda,
swamp drainage for rice and sugar cane plantations threa-
tens the Shoebill.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 305.
 American
White Ibis
Eudocimus ruber (Linnaeus)

Scolopax rubra Linnaeus, 1758, syst. Nat., ed. 10, p. 145; based chiefly on 'The Red Curlew' of Catesby,
1732, Nat. Hist. Carolina, pt. 5, p. 84, pi. 84: America ( = Bahamas)

Subspecies:
Eudocimus ruber ruber (Linnaeus)
Eudocimus ruber albus (Linnaeus)
Scolopax alba Linnaeus, 1758, Syst. Nat., ed. 10, p. 145; based on 'The White Curlew' of Catesby,
1732, Nat. Hist. Carolina, pt. 5., p. 82, pi. 82: America (=Carolina)

Other names:
Scarlet Ibis, White Curlew, Spanish Curlew, Red Curlew, Lesser Ibis, Currie Currie, Brown Ibis
[juvenile], Brown Curlew [juvenile] (English); Corocoro, Corocoro Rojo, Corocoro Blanco, Corocoro
Colorado, Coco, Coco Blanco, Curri-curri, Ibis Blanco, Ibis Escarlatea, Coclito Blanco, Coclito
Colorado, Coclito Rojo, Coclito Mojoso [juvenile] (Spanish); Flamant, Ibis Flamant, Courly Rouge
de Bresil, Ibis Rouge, Ibis Blanc (French); Rode Ibissen, Rode Ibis, Witte Ibis, Scharlaken Ibissen,
Korikori, Flamingo (Dutch); Guara (Portuguese).

IDENTIFICATION
The American White Ibis, more appropriately called the Scarlet Ibis in South America, is easily
identified as a medium-sized ibis (56-68 cm long) typically with pure white or red plumage. Over most
of its range, it is often the most abundant wading bird and, because of its numbers, flocking habits and
obvious plumage, one of the more spectacular.
The basic plumage of the adult is a solid colour, an immaculate white, a deep scarlet-red or an
orange-pink. The wing-tips are glossy blue-black, so dark as to appear black in the wild. The facial skin,
bill and legs are pink-red, and the iris is blue; darker in scarlet birds. The species lacks specialized
plumes such as are found on many other species of ibis.
The reddish plumage is variable among individuals and populations, in that the colour depends on a
combination of season, food and genetics. In northern South America, individuals of various shades of
pink or orange typically occur among the darker red birds. Birds are duller and more variably coloured
in the inland marshes of Venezuela than on the coast (Luthin 1983a, P.O. Frederick pers. comm. 1990).
Some mostly white birds have pink mottling.
Males and females have identical plumages, but the male is substantially larger (Kushlan 1977d).
Perhaps this difference is most apparent in bill length: the male's bill is 1.24 times that of the female in
both subspecies (Kushlan 1977d, Luthin 1983a). The differences are such that they can be used to
identify the sexes in the field, especially when they are together.
During courtship, the bare skin of the legs, face and upper bill turns bright red, primarily through
increased vascularization (Frederick 1987c).
The bill usually darkens during courtship, but this feature is variable in ways that are not entirely
148 American White Ibis

comm.) speaks of seeing a small compact sac on either side

of the throat rather than one sac. The photography of Ramo
and Busto (no date) clearly show development is more
extensive in the female Scarlet Ibis. Additional data are
needed regarding geographic, sexual and individual differ-
ences in throat-sac development.
Colour changes are very fleeting, and the intense colours
fade soon after courtship is over. The intense skin colour
pales almost immediately. Bills begin to fade after a few
weeks, although a dusky wash can remain for months.
During nesting, white plumage becomes soiled and disco-
loured and such birds can appear to have a dark or buffy
wash. In the llanos, this wash may be yellowish (Ramo and
Busto no date:81).
A complete post-nuptial moult begins as nesting winds
down and is well underway by the time nesting is com-
pleted. This moult begins on the neck and back, spreading
to the wing scapulars, coverts, tail and finally the primaries.
In the red form, necks may turn a pale pink as a result of
this moult (M.v. Wieringen and K. Brouwer pers. comm.
1988), making the birds look scruffier and duller than in
breeding plumage, and accentuating individual differences.
An incomplete moult occurs prior to nesting (Beebe 1914).
The chick appears grey. The head, back and wings of
newly hatched young are covered with a grey to dark black
down, more blackish on the head and more brownish on the
wings. The ventrum is initially bare, becoming white-
feathered. Some individuals have a patch of white feathers
on the head. This distinctive tuft may persist throughout
nesting and provides a measure of individuality (De Santo
et al. 1990). The skin is initially pink but turns grey within
the first 2 days of hatching; the iris is brown; the feet and
documented as yet. In general, the distal portion of the bill legs are pink-grey but turn grey during the nestling period
darkens to a glossy black, and this darkening extends (Beebe 1914, ffrench and Haverschmidt 1970, Antas 1979,
towards the head to a variable degree. In North America M.v. Wieringen and K. Brouwer pers. comm. 1988, De
usually about two-thirds of the bill darkens, whereas in Santo et al. 1990). The short, comparatively straight bill of
some individuals the colouring is more extensive. In north- the hatchling is pink with a dark tip, but becomes ringed by
ern South America, the entire bill is usually coloured in both a variable number of black bands (usually 3), starting to
white and red birds, but many have only partial darkening develop at c. 6 days after hatching (M.v. Wieringen and K.
(see Ramo and Busto, no date). In southeastern South Brouwer pers. comm. 1988). The coverage and pattern of
America, the female's bill remains brownish with a blackish the rings vary among chicks. By 6 weeks, the entire bill is
tip during the breeding season (H. Sick 1985, pers. comm. grey and then turns pinkish over the next few weeks (Beebe
1990). The pattern of variability in this seasonal feature 1914, De Santo et al. 1990).
requires much additional study before its geographic trend Juvenile plumage, which differs from that of the adult,
can be understood. begins to develop as the primaries emerge on the fourth or
Also during the height of pair-formation, courting birds fifth day (De Santo et al. 1990). Other wing and body
develop a bright red, naked throat-pouch, which can extend feather tracks emerge within 2 weeks, and the plumage is
up to 3 cm. Both sexes develop some expansion of the bare generally complete by the end of the third to fourth week
throat area, but the extent of development is variable among except for retention of down on the head and neck through
individuals and populations (Beebe 1914, Kushlan 1973b, the fifth to sixth week. The head, back, upper breast, wings
Rudegeair 1975a).Jn North America, the female usually and tail are slate grey and the undersides white; the face and
(Kushlan 1973a, Rudegeair 1975a) but not always (Beebe bill are pink, the legs grey and the iris slate grey.
1914, C.R. Schmidt 1968) seems to have a more developed Juveniles fledge at a smaller than adult size and continue
sac. It has been reported that in South America both sexes to grow after fledging, taking up to 2 years to achieve adult
develop a sac (Luthin 1990, Wieringen and Brouwer 1990). weight (Kushlan 1977f). Beginning as early as 4.5 months
However, the primary source of this information, Risdon old in captivity and certainly by 8-9 months old in the wild,
(1969), described wattles on both sexes, rather than a gular the juvenile feathers, now faded to a sandy grey, begin to be
sac, and that author admitted to being unable to tell the replaced with white or red feathers resulting in a mottled
sexes apart. Sick (1985, translated by Belton 1990, pers. appearance until moult is completed. Most of the plumage

has changed by the end of the second year, but brown
mottling remains on the head and neck into the third year.
First nesting takes place when young are 3 years old.
Vocalizations are variable, but are generally character-
ized as honks. The primary flight and disturbance call is a
coarse 'Hunk-hunk-hunk'. Nestlings beg with a distinctively
shrill 'Zziu'. They often vocalize while feeding, the flock
giving off a murmuring, muffled 'Huu-huu-huu'. During
courtship 'Hunking5 and squealing calls are given.
The American White Ibis is an accomplished aerialist,
the birds flying in excess of 40 km/h, alternating rapid wing
beats with gliding, typically in a variable V-formation (Pen-
nycuick and De Santo 1989, pers. obs. J.A.K.). They soar to
great heights. This behaviour is especially spectacular over
a colony, even before the initiation of nesting. The birds also
soar when gaining height to glide to feeding sites. Upon
disturbance, a foraging aggregation will take to flight all at
once and may perform manoeuvres before returning to the
feeding site or landing on a roost nearby to await passage of
the disturbance. Individual birds may engage in spectacular
aerobatics. Especially characteristic is a free fall into a roost.
Young and adults tend to flock and feed separately; typi-
cally first- and second-year birds are found together.
DISTRIBUTION AND POPULATION
Overall, the range of this species is quite extensive, but the
breeding range is more patchy. It nests over much of the
coastal plain of southeastern North America through Cen-
tral America, the Greater Antilles, in northern South Amer-
ica, and in southeastern Brazil.
The subspecies albus occurs in the northern portion of the
range through Central America and perhaps into western
South America. Although the line of demarcation between
races is not clear, the subspecies ruber is found in the remain-
ing part of the South American range.
In North America, the American White Ibis has been
slowly expanding and consolidating its range for the past
several decades and now nests as far north as Virginia,
south through Florida, to Louisiana (Sprunt 1944, Frohring
and Beck 1978, Shields and Parnell 1983). It also nests in
Baja California, both coasts of Mexico, Belize, both coasts of
Nicaragua (Chinandega, Leon, Managua, Chontales, Car-
ibbean Region of Zelaya), the Pacific but not the Caribbean
coast of Costa Rica (Guanacaste, Puntarenas including
Golfo de Nicoya and Golfo Dulce), and Panama (Panama
Bay, Herrera, Los Santos, Chiriqui) (Slud 1964, Luthin
1984a, D.A. Scott and M. Carbonell 1986, Stiles pers.
comm. 1988, Tejera 1989a,b). In the Greater Antilles, it
nests in Cuba, Hispaniola and reportedly in Jamaica (Gar-
rido and Garcia 1975, D.A. Scott and M. Carbonell 1986).
In South America, nesting occurs in northwestern Colom-
bia, interior and coastal Venezuela, and along the Atlantic
coast through Guyana, Surinam, French Guiana and north-
eastern Brazil in Amapa to Maranhao, and in southeastern
Brazil (Sick 1985, D.A. Scott and M. Carbonell 1986,
Marcondes-Machado and Monteiro 1989, 1990, Teixeira et
al. 1990, Antas et al. 1990). It previously nested in Trini-
dad, along the northeast Brazilian coast to Ceara and along
American White Ibis 149
the southeast Brazilian coast (from Rio de Janeiro to Santa
Catarina).
The relative abundance of white and red birds varies over
the overall range. Northern breeding populations are com-
posed primarily of white birds, but not entirely so as scarlet
birds have regularly been reported from the range of white
birds since their description from there by Catesby two
centuries ago. Fully scarlet birds continue to nest in the wild
in Florida (pers. obs. J.A.K., P. Frohring pers. comm.
1990). The origin of these birds must remain uncertain
owing to previous introductions, but the fact that we have
observed them regularly for two decades is of some import-
ance. In South America, nesting ibises are mostly red birds,
with far fewer white birds from northern Colombia through
Venezuela to Surinam. Red birds also are typically found
down the Pacific coast to Ecuador and northeastern Peru
(F.P. Penard and A.P. Penard 1908, Ramo and Busto 1982,
1984, 1988, Luthin 1984a, Serna 1984, Hilty and Brown
1986). In the llanos, about 4-6% (depending on where they
are counted) of the birds are white (Ramo and Busto 1987).
This highly nomadic species undertakes seasonal mi-
grations, intra-regional population shifts and wide ranging
dispersals (Palmer 1962, Kushlan 1976d, 1979c, Kushlan et
al. 1985). In migration, North American birds move into
Florida and Mexico, and perhaps into Cuba and Central
America. This migration mostly goes around the Gulf of
Mexico, with post-migration arrival dates at the northern
Gulf coast in first 2 weeks of March (Stevenson 1957).
Population shifts on a similar scale also occur. In Florida,
the population shifts between seasonal interior wetlands and
the coast (Kushlan 1976d, 1979c). A similar shift occurs in
Venezuela, from the higher llanos to middle and lower
llanos and to the coast (in Dec-May) where birds are also
found year-round (Luthin 1983a, M.v. Wieringen and K.
Brouwer pers. comm. 1988, McNeil et al. 1990, Kushlan et
al. 1985). Coastal nesting birds also migrate, as evidenced
by their seasonal occurrence in Trinidad (Hislop and James
1990). It has long been thought that these birds nest at the
mouth of the Orinoco in Venezuela (fFrench 1967). Scarlet
birds breeding in Surinam shift along the coast after the
nesting (Held 1990). Shifts also take place in Brazil with
birds sometimes moving up the Amazon into Para (Sick
1985) and to the coast in Sao Paulo and Parana (Guiz pers.
comm. 1990).
Wide-ranging dispersals take white and scarlet birds out-
side of their usual nesting and migratory ranges, and in fact
they have been reported as vagrants or rare visitors over
much of North, Central and South America. The extent of
dispersal varies among years, probably related to water and
feeding conditions. In North America, the species occurs
occasionally as far as New York, Quebec, Pennsylvania,
Illinois, Missouri, Kentucky, Wyoming, South Dakota, Col-
orado and California (e.g. Dresser 1866, Lincoln 1923,
Palmer 1962, D.H. Davis 1979, Ernst 1974, Irmscher and
Irmscher 1979, J.C. Miller 1982a,b,). White birds also
occur rarely along the Caribbean slope of Nicaragua and
Costa Rica (G. Stiles pers. comm. 1988), in the Bahamas,
Jamaica, Cayman Islands, Puerto Rico, Trinidad, Aruba,
Curasao, and along the Pacific coast as far south as Tumbes
150 American White Ibis
in northwestern Peru (Brudenell-Bruce 1975, Schulenberg
and Parker 1981, Raffaele 1983, Bradley 1985).
Red-plumaged birds are naturally accidental in Jamaica,
Grenada, Cuba and elsewhere in the West Indies, and also
(despite views to the contrary) in North America (Audubon
1840, Brewster 1883, Sell 1918, R.T. Dusi 1965, Simpson
1966, Cypert 1971, Garrido and Garcia 1975, Maehr and
Hintermister 1982). A record from Honduras is considered
to be doubtful (Monroe 1968). Scarlet birds also have been
seen east of the Andes in Ecuador (Stevenson 1972).
Population sizes of white and scarlet ibises are among the
best known of any ibis species, because they nest in large
colonies and are historically the most numerous ciconiiform
wading birds in most of the areas in which they occur (with
the possible exception in recent years of the immigrant
Cattle Egret, Bubulcus ibis).
Along eastern North America, over 64000 were counted
nesting in about 60 coastal colonies in 1975 (Custer et al.
1980). Over 87000 nested in Louisiana in 1967 (Portnoy
1981). Colonies of over 10000-20000 birds have recently
been reported from coastal North America (Osborn and
Custer 1978, Frederick 1986a, Bildstein et al. 1990b). How-
ever, the largest of these colonies have been abandoned in
the past few years (W. Post 1990, K.L. Bildstein pers.
comm. 1990).
In southern Florida, historical populations numbered
near 100000; reports of historic numbers in the millions
have no credence (Frohring et al. 1988). Until very recently,
populations have remained relatively high there; in the
1970s populations numbered over 50000, including over
35000 in one colony (Kushlan 1973a, Kushlan and White
1977). Nesting numbers have been decreasing rapidly in the
Everglades in recent years.
In Mexico, numerous small colonies of 20-600 nests occur
along the Pacific coast and smaller numbers of larger colon-
ies, of more than 2000 nests, have been counted on surveys
on the Gulf of Mexico coast (Knoder et al. 1980). More
recently, 1750 pairs were counted nesting at Bahia de Santa
Maria in Sinaloa, 1250 pairs in Tabasco and 1000-1500
pairs in Quintana Roo (D.A. Scott and M. Carbonell
1986).
In Central America, colonies number in the hundreds to
thousands. In Belize, numbers may be increasing; 400-500
pairs recently nested in Belize District (D.A. Scott and M.
Carbonell 1986, D.S. Wood and R.C. Leberman 1987).
Colonies are found on the Pacific coast of Costa Rica and in
Panama; about 1000 pairs nest at Palo Verde and 600 near
Canas in Guanacaste, Costa Rica (Delgado in Luthin
1984a, D.A. Scott and M. Carbonell 1986, Stiles pers.
comm. 1988).
In northern South America, the scarlet subspecies of the
American White Ibis remain abundant and recent surveys
have well-documented its nesting status (Spaans 1975a,
1982b, Ramo and Busto 1984, 1988, Luthin 1984a,
Dujardin and Tostain 1985, D.A. Scott and M. Carbonell
1986, C.S. Luthin pers. comm. 1990, C. Ramo pers. comm.
1990, Ramsamujh 1990, Held 1990, Dujardin 1990, Hislop
and James 1990, Teixeira et al. 1990, Antas et al. 1990).
Numbers in Colombia are poorly known, but birds nest
along the coast and in the western llanos of that country.
The population numbers in the hundreds of thousands in
northeastern South America. The total in Venezuela is over
128000, including a colony of over 52000 birds; in the
Venezuelan llanos, over 95% of the nesting population
surveyed in 1983 were found in 22 colonies (Ramo and
Busto 1988).
About 300 pairs are known to have nested in Guyana
in mid 1970s (Ramsamujh 1990). Nesting populations in
Surinam numbered over 13000 pairs in the early 1970s,
decreasing over the decade to 2000 nesting pairs, then
increasing to nearly 35000 pairs in 1986 (Spaans 1982b,
K. Brouwer pers. comm. 1988, Held 1990). In French
Guiana, populations totalled about 3000 pairs in 1988
(Dujardin 1990). In 1991, 1700 pairs were counted in two
colonies (Dujardin 1991). In Trinidad, ibises have been
counted at their roost-site for many years (Hislop and James
1990). Birds using the roost increased from 1500 in 1969 to
10000 in 1987, but no nesting has taken place in Trinidad
for 30 years.
In Brazil, the species' status is becoming better under-
stood (Luthin 1984a, Sick 1985, Bokermann and Guix 1987,
1990, Teixeira et al. 1990, Antas et al. 1990), although
censuses of nesting colonies are few. In the northeast, one
colony included approximately 5000 ibis nests. The species
was once reported to be the most common wading bird of
the Amazon region but over the last century has become
scarce there (Sick 1985). In 1981-82, 7500 adults were
counted at the mount of the Amazon. During the non-
breeding season, up to 9000 have been counted by R.G.
Morrison (pers. comm.) in the northeast of Brazil. In the
southeast of Brazil, these ibises were thought to have been
extirpated, but birds have been observed there repeatedly
since the early 1980s, particularly in Rio de Janeiro and Sao
Paulo states. A small nesting colony was recently discovered
at the junction of the Oncas and Quilombo Rivers
(Marcondes-Machado and Monteiro 1990).
Throughout the range of this species, because of its pro-
pensity to change colony sites, nesting numbers fluctuate
from year to year and decade to decade in any one area, and
such fluctuations may or may not reflect the underlying
population sizes (ffrench and Haverschmidt 1970, Kushlan
1979b, Spaans and de Jong 1982, Dujardin and Tostain
1985, M.v. Wieringen and K. Brouwer pers. comm. 1988,
W. Post 1990). For example, along the eastern North Amer-
ican coast, nesting numbers decreased 40% within 1 year,
between 1975 and 1976 (Custer et al. 1980). Similarly, at
two colony sites in North Carolina, nesting numbers de-
creased from 12 000 pairs at each site in the wet year of 1984
to 2000 pairs at each in the following very dry year (Bild-
stein et al. 1990a). Population fluctuations along the Suri-
name coast have been similarly extensive and impermanent.
Some population fluctuations have resulted in local long-
term decreases, however. This is especially the case in
southern Florida, the Atlantic Coast of North America,
French Guiana and southeast Brazil.
The nomadic tendency of this species also suggests a low
degree of isolation among populations. This has been shown
to be the case genetically (Stangal et al. 1991).

ECOLOGY
The American White Ibis is a bird of interior and coastal
marshes and swamps. Historically it probably reached its
peak abundances in the great grass and sedge wetlands of
the eastern USA and Venezuela, but throughout most of its
range it occurs in coastal areas. In southeastern Brazil its
historic range coincided with the extent of mangrove
swamps. However in North America, it extends well into
temperate areas, where it uses salt marsh and nearby fresh-
water swamps.
This ibis actually can use a wide variety of feeding habi-
tats, usually in shallow water (frrench and Haverschmidt
1970, Kushlan 1979c, Guster and Osborn 1978, Desenne
and Shimotake 1990). Inland, it is most typically found in
shallow seasonal marshes, swamps and river-edge hard-
wood forests, and almost as often in wet pasture and shallow
ephemeral ponds. Along the coast its typical feeding habitat
is marsh and mangrove swamp, where feeding is dependent
on tides (Desenne and Shimotake 1990).
Evidence has accumulated that the American White Ibis
suffers reduced nesting success when restricted to foraging
in salty habitats. So they often choose freshwater feeding
habitats, even if the nesting colony itself is situated in a
marine or estuarine environment. Several studies have
found smaller clutch sizes on the coast than inland (Kush-
lan 1977d, Rudegeair 1975b). More recent studies have
shown that young are incapable of enduring the high salt
content of marine food (J.W.Johnston and K.L. Bildstein,
1990).
Throughout its range, the species can take advantage of
particularly good feeding opportunities that happen to be-
come available. It should not be unexpected to see this
species feeding in such places as open marine lagoons, on
exposed tidal flats, along the tide line on beaches or even in
the surf, in gardens, residential yards, ball fields and golf
courses, in agricultural fields, following tractors, in garbage
dumps, or in the accumulating manure under chicken
coops. It feeds heavily in rice fields at the beginning of the
wet season in the llanos (Wieringen and Brouwer 1990).
Ibis populations move about to take advantage of avail-
able foraging opportunities, establishing a succession of
roosts as conditions change from year to year, or even as a
season progresses. Feeding locations are usually within
10 km of a colony site, although we have followed flights for
over 40 km in southern Florida, as has Bildstein in coastal
Carolina (Bateman 1970, Kushlan 1976d, 1979c, Custer
and Osborn 1978, K.L. Bildstein pers. comm. 1989).
This species can feed effectively whether water levels rise
or fall as long as foraging sites of appropriate water depth
are available (Kushlan 1979c). In general it appears that
water depth is more important than direction of water-level
change in determining foraging patterns of ibises in the
Florida Everglades.
A flocking species, American White Ibises may feed in
large aggregations and fly in cohesive flocks between feeding
and roosting sites (Kushlan 1978a, Wrege 1980, Petit and
Bildstein 1987). Juveniles often flock separately, taking
2 months to begin flock flying; they are less successful at
American White Ibis 151
foraging than are adults and may feed on the periphery of
adult flocks (Bildstein 1983, 1984, Petit and Bildstein 1986).
Feeding aggregations form by local enhancement—the
attraction of birds to other birds already feeding at a site,
which is one way birds are able to locate suitable feeding
locations (Kushlan 1976b, 1978a, 1981). Birds foraging
within a flock have an advantage in being able to spend less
time on surveillance (Petit and Bildstein 1987).
The American White Ibises feed primarily by non-visual
Probing in shallow water but also use such techniques as
Head Swinging, Groping and Pecking when appropriate
(Kushlan 1977c). They hunt crabs visually. Their usual
behaviour is to Probe deeply in the water while Walking
Slowly. They may Stand to concentrate their Probing under
a plant or within a crayfish or crab burrow. On dry or
mostly dry land they generally feed visually, picking up prey
items from the surface of the soil (Pecking). Details of the
foraging techniques, such as rates of probing, capture, walk-
ing and turning, vary depending on the nature and avail-
ability of the food supply (Henderson 1981).
The close association with other wading birds can result
in intense interactions among individuals. Ibises defend
their individual distances aggressively and are subject to
having their captured prey stolen by nearby birds (Kushlan
1978c, Kilham 1984). Ibises can be aggressive and can
successfully steal prey from other species (P.C. Frederick
pers. comm. 1990). Conversely, they are used as 'beaters' by
other species that walk after them as they forage (Courser
and Dinsmore 1975, Kushlan 1978b, Kilham 1980, 1984).
Desenne and Shimotake (1990) found them feeding com-
mensally with capuchin monkeys (Cebus sp.).
The diet has been well documented. The American White
Ibis is a specialist on aquatic invertebrates, with the bulk of
its diet in most instances consisting of crayfish, small crabs,
and aquatic insects and their larvae. Their feeding methods
make them selective feeders on these prey in shallow fresh-
water habitats (Kushlan 1979b,c). In coastal marshes, they
seek out fiddler crabs (Uca spp.), the females of which
appear to be more vulnerable to predation (Bildstein et al.
1989). All types of aquatic and terrestrial insects are taken,
along with a wide variety of other prey types (Nesbitt et al.
1974, Kushlan and Kushlan 1975, Haverschmidt 1968, Sick
1985, Kushlan 1979c, M.v. Wieringen and K. Brouwer
pers. comm. 1988, 1990, Tejera Nunez 1990, Desenne and
Shimotake 1990). Among their other prey, they eat worms,
snails, bivalves, frogs and small fishes, when they can catch
them, and almost any other small item they can grab and
handle.
Adults require 165kcal/day; young require 8620 kcal of
food during the nestling period (Kushlan 1977b,c). The
larger male requires more energy than does the female, and
the male's size is detrimental to meeting his energy require-
ments because he obtains no compensating increase in for-
aging effectiveness (Bildstein 1987).
Because of its large population sizes, the American White
Ibis can contribute substantially to energy flow through the
avian community. The southern Florida population of
62000 birds required 930 000 000 kcal in 1 year (Kushlan
1977b,c). In a temperate saltmarsh, the species accounted
152 American White Ibis
for 17% of the avian biomass and 10% of the energy flow
through the bird community of the wetland (Christy et al.
1981; Bildstein et al. 1982a, K.L. Bildstein pers. comm.
1989).
BREEDING
In the American White Ibis breeding is highly communal,
colonies often being both large and densely packed with
nesting birds. This species usually nests in trees and bushes
but also nests in such places as cactus, grasses and sedges,
or on the ground. Colonies are situated especially on true
islands, on islands of trees within marshes, in mangrove
swamps and in gallery forests. Except on true islands, the
presence of water under the nest sites seems to be crucial to
colony formation. Colony sites usually develop at roost sites
through the formation of daytime bachelor parties of males
that remain at the roost and on the nearby ground to begin
their display activity.
Throughout its range, the species is highly vagile in its
use of nesting colony sites. Populations move around fre-
quently, and it is more typical for a site to be used only once
or just a few times than for it to be used regularly over many
years. This is clearly a rather nomadic species.
In general, nesting appears to take place when local
conditions are suitable for effective foraging, primarily the
availability of sufficiently shallow wetlands or the shallow
flooding of higher habitat (Kushlan 1976d, 1979b). In the
Florida Everglades, this is usually during drying conditions,
although ibises there can feed effectively on both falling and
rising water levels (Kushlan 1979c). Along the coast of
North America and in the llanos, nesting is on rising water
levels at the onset of the wet season (M.v. Wieringen and K.
Brouwer pers. comm. 1988, K.L. Bildstein pers. comm.
1990, C.S. Luthin pers. comm. 1990). Nesting along the
coast of South America is more variable; and, although they
appear to be less influenced by climatic conditions, nesting
is mostly in the wet season (M.v. Wieringen and K.L.
Brouwer pers. comm. 1988, C.S. Luthin pers. comm. 1990).
Because of their dependence on water conditions, the
actual timing of nesting varies throughout the species' wide
range, and also in any area from one year to the next. In
South Florida, American White Ibises may nest any time
from the more usual spring nesting period (Mar-Apr),
through the summer, and even into the autumn (Rudegeair
1975a, Kushlan 1976d, Allen-Grimes 1982). Nesting may be
a bit later in coastal colonies, lasting into the summer.
Timing in coastal Carolinas is similar, April-May (K.L.
Bildstein pers. comm. 1990). In the llanos, they begin to
nest in June and July (M.v. Wieringen and K. Brouwer
pers. comm. 1988, Luthin 1990). In Trinidad, egg-laying
varies from April to September (ffrench and Haverschmidt
1970). Nests with eggs have been recorded in Surinam from
April to August, and in French Guiana in mid-April to June
(Haverschmidt 1968, Dujardin 1990). Less exact infor-
mation is available from Brazil; in the northeast, active
colonies were found from March to September (Antas et al.
1990).
Courtship behaviour has been well documented, and
there appears to be no substantive differences between the
colour forms (Palmer 1962, Kushlan 1973b, 1977d, Rude-
geair 1975b, Spil et al. 1985, Ramo and Busto 1985, C.S.
Luthin pers. comm. 1990). Display Sleeping, Display
Shaking, Display Preening and Supplanting Flights are
common displays seen in bachelor parties. Display sites are
tightly defended by males with Forward display and short
Supplanting Flights. Display Preening, Head Rubbing and
Bill Popping are the primary Displays of the male. Head
Rubbing is quickly initiated and may include a complete
roll of the bill from left to right or only a quick touch of the
back of the head in a pumping motion, rather reminiscent of
the Stretch display of herons. Full Bill Popping includes
twig grasping and nibbling, and may be accompanied by
complete extension of the neck until the head is down
between the legs. In South American birds a rocking motion
has been seen (M.v. Wieringen and K. Brouwer pers.
comm. 1990).
A female approaches displaying males very cautiously in
the Bowing display, especially showing the side of her face
to the male. This posture presents the fire red face and gular
pouch, highlighting the eye (Kushlan 1977d). The male
gives Forward display and may actually attack, sometimes
going so far as to grab the female's head and shaking it like a
twig; this often appears to be redirected Twig-grasping.
M.v. Wieringen and K. Brouwer (pers. comm. 1988) did
not see such head-grasping in South America. Before pair-
bonding is completed, the female can become quite bloodied
in her attempts to remain in the male's display area.
During courtship, the pair uses Greeting displays and
mutual Stick Shaking, typically with necks crossed. Both
also spend considerable time Display Preening and Stand-
ing touching each other. Greeting displays may include
twig-grasping and hunking vocalizations by the males and
squealing vocalizations by the females, the latter ceasing
later in courtship.
Nests are usually near but not necessarily on the display
site; actual nest-site selection falls to the female, who builds
the nest with sticks brought by the male. She shoves the twig
into the nest-platform and positions it by moving it up and
down in a tremble shoving motion. Nests are typically
closely grouped, even almost touching each other. Nest-
platforms often contain fresh leaves, replenished by the male
during incubation.
Copulation begins with the male placing his neck over
that of the female as they engage in mutual twig-grasping.
The male then mounts, achieving and maintaining his posi-
tion by treading his feet and flapping his wings. Copulations
occur throughout the day.
The first egg is laid 5-6 days after the beginning of
copulations, and additional eggs are laid at 1- to 2-day
intervals, the latter being usual (P.C. Frederick pers. comm.
1990). Extrapair copulations are common in the later stages
of pair-formation and even into incubation. The male
initiates these—the female, although capable of resisting,
seldom does so—and it is estimated that over 6% of success-
ful copulations are outside of a mated pair (Frederick
1987a).
Clutch size is 2-5, averaging 2-3 depending on location:

clutch sizes averaged 1.97 in coastal Louisiana, 2.07 in
inland northern Florida, 2.20 on the Florida coast, 2.23 in
coastal North Carolina, 2.43 in Venezuela, 2.45 in the
Everglades, 2.49 in interior central Florida (Kushlan 1977c,
Rudegeair 1975b, Allen-Grimes 1982, Shields 1985, Ham-
matt 1981, B. Busto and C. Ramo pers. comm. 1988). Eggs
are smooth and not glossy, splotched with brown over a
base colour ranging from cream to blue-green. Shape varies
from ovate to elongate, some surprisingly elongate.
Nest-attendance patterns of the parents depend on the
stage of nesting and distance to the feeding sites (Kushlan
1976c, M.v. Wieringen and K. Brouwer pers. comm. 1988).
The male tends to remain at the nest throughout the nest
building period, as departures put the nest and its contents
at considerable risk (Kushlan 1973b, Frederick 1986a).
During the 21-day incubation period and during early
hatching, the pair alternates nest attendance. In the Ever-
glades, the males were observed to remain at the nest during
the day early in nesting (Kushlan 1973b), but this probably
is more related to food location and the number of foraging
trips. Observations in other locations and later in nesting
indicate that the parents alternate more frequently (Wier-
ingen and Brouwer 1990), and this is probably the more
typical situation.
It is clear that the larger size of the male is effective in
defending against nest predation or nest material robbing,
in that lone females are susceptible to being driven from the
nest site (Frederick 1986a). Contests between males seem to
be settled by size difference (P.C. Frederick pers. comm.
1990). Predation by Black-crowned Night-Herons (Nyeti-
corax nyeticorax), crows (Corvidae), and other birds is a
particular threat to unguarded young (Kale 1965, Shields
and Parnell 1986). Crows may steal eggs and unintentio-
nally drop them in other nests (Shields 1987). The guardian
defends the nest or young with Forward display and Bill
Thrusts. Renesting after disturbance is possible, but its
frequency is not known.
Eggs in a clutch hatch 1-2 days apart, and the hatchlings
are incapable of even holding their head up for the first day
or so. To feed the hatchling, the adult grasps the young's bill
stimulating it to raise its head and then regurgitates into its
mouth. After a week, the young initiate feeding with a
Begging display by touching the parents bill while calling
and vigorously flapping their wings. Males may pirate food
from females and young, and considerable piracy occurs
among juveniles (Frederick 1987a, De Santo et al. 1990).
The feet of the young develop rapidly and the birds can
move from their nest as early as 2 weeks, forming creches
(Kushlan 1977f, K. Brouwer and M. van Wieringen 1990a).
At this time, both adults are away from the nest foraging at
the same time, returning to feed the young up to 9 times/
day. Young leave the colony when about 50 days old, at
which time the bill is under-developed relative to adult size
whereas the legs are nearly fully grown, an adaptation for
walking foraging (Kushlan 1977f, De Santo et al. 1990).
Nesting success is often poor, usually averaging near one
or so young per nest and about a 50% mortality (Kushlan
1977c, Girard and Taylor 1979, Allen-Grimes 1982, Freder-
ick 1987a). Colonies suffer large mortality due to high tides,
American White Ibis 153
predation or lack of food. In southern Florida, colonies
fledged about 1.03 young per nest. In Louisiana, 64% of all
nests were lost during incubation by high tides (Hammatt
1981). A similar situation in South Carolina led to repeated
flooding at a coastal colony, and subsequently its total
elimination by a hurricane (A. Dinep pers. comm. 1990,
K.L. Bildstein pers. comm. 1990). In this site, production of
fledglings decreased from 39% in a wet year to 3% in a dry
year (Bildstein et al. 1990a). Over its range, rainfall and
water-level conditions are usually the primary determinant
of nesting success in this species.
TAXONOMY
The Eudocimus ibises (Parkes 1951) appear to be most closely
related to the Plegadis ibises (Mayr and Short 1970). How-
ever, the Plegadis ibises have a distinctive conformation of
the tarsometatarsus and so are readily distinguishable
(Olson 1981). Eudocimus is the more primitive group, with
the fossil Plegadis paganus being somewhat intermediate.
Examination of additional fossil material would be desirable
to resolve the question of derivation of the Plegadis ibises
from Eudocimus stock.
The relationship between extant North and South Ameri-
can populations of the genus Eudocimus has long been a
matter of interesting discussion. Ridgway (1884) long ago
proposed that they were the same species. However, sub-
sequent interpretations gave special credence to one of the
earliest visits to a colony in the llanos by Zahl (1950), who
found no mixed-colour pairs, although his visit was very
short. None the less until recently, this weak evidence
suggesting that the two forms do not interbreed has
influenced interpretations of their taxonomic relationships.
Recent studies, especially by Cristina Ramo, Benjamin
Busto, Charlie Luthin, Eduardo Aguilera Prieto, Koen
Brouwer, and Marcel van Wieringen, have provided addi-
tional evidence. Ramo and Busto (1982) formally proposed
that the white and scarlet ibises be considered to subspecies.
As first revisors, they chose the name Eudocimus ruber for the
species.
The primary differences of possible taxonomic import-
ance between North and South American populations are in
plumage colour, size and some details of the soft-part colo-
ration. The fundamental taxonomic question is: how differ-
ent are these forms and are these differences sufficient to
inhibit interbreeding if they were in geographic contact?
The most obvious distinction, of course, is in plumage
coloration of the adult. (The juveniles are identical.) The
red coloration of the birds is due to the deposition of specific
carotenoid compounds secondarily derived from dietary
pigments (Fox 1962, Fox and Hopkins 1966). This implies a
genetic basis. However, the actual colour expressed in an
individual (its phenotype) is the result of a combination of
dietary influences and genetics. As a result, plumage colour
is variable within and between populations. In the llanos,
for example, birds range from deep scarlet, to orange, to
white.
The genetic component is probably a simple difference in
a biochemical pathway that could be under the control of a
154 American White Ibis
single enzyme system, and perhaps a single gene. Salmon-
coloured birds result from the mating of white and red
adults, the faded colour apparently resulting from the in-
complete dominance of the dominant colour gene. If this
indeed is the case, back-crossing of a genetically salmon-
coloured bird (heterozygote) to a homozygous white or red
bird would produce either another pink bird or a pure white
or red bird like the parent. Thus, we would expect to see all
three forms in the population, which is the case in South
America. That the genetics is slightly more complicated is
suggested by the occurrence of slightly mottled white and
pink birds.
South American birds, both white and red, are smaller
than those from North America. However, such a size cline
is to be expected in a bird which ranges from the temperate
to tropical climates. Wieringen and Brouwer (1990) provide
strong evidence for the existance of such a cline in their
comparison of the sizes of birds from Florida, Venezuela
and Surinam. There remains more differences in size be-
tween the sexes than between the two colour forms, how-
ever.
It has been suggested that the North and South American
forms differ in soft-part colours in that the entire bill of the
scarlet ibis turns black during the height of courtship in the
South American birds whereas it only partially turns black
in North American birds. However, there is much variation
and these colours hold for only a very short period at the
height of courtship. Some South American birds do not have
a fully black bill (see photos in Ramo and Busto no date)
and some North American birds do (pers. obs. J.A.K.). H.
Sick (1985, pers. comm. 1990) stated that, in the southern
South American populations, the bill of the male turns shiny
black but the female's bill stays brownish with a blackish tip
only; this also occurs in northern South America.
The presence of the throat-sac is another character of
interest. A case has been made that in North America the
female ibis has a throat-sac during breeding, whereas both
sexes have a sac in South America (Wieringen and Brouwer
1990). However, both sexes show some enlargement of the
throat during courtship in both populations (Beebe 1914,
C.R. Schmidt 1968, pers. obs. J.A.K.). Several observations
do suggest that more males may have obviously enlarged
sacs in South America, although the citation generally used
to support this argument (Risdon 1969) notes the occur-
rence of small wattles, not fully developed throat-sacs (he
also admitted that he could not tell the sexes apart).
All observations indicate that the courtship behaviour of
the two forms are virtually identical (Spil et al. 1985, C.S.
Luthin pers. comm. 1990). Some minor differences have
been reported under zoo conditions, such as a scarlet ibis
rocking with its head down (M.v. Wieringen and K.
Brouwer pers. comm. 1988). But, of course, birds used to
each other, especially in captivity, could be expected to
exhibit somewhat artificial behaviour.
If one were to draw a conclusion from the above dis-
cussion alone, the weight of the evidence provides no reason
why these two clearly related forms should not be con-
sidered the same species. However, this is inferential and
definitive evidence can only be obtained from observations
of interbreeding. Fortunately such evidence exists, and it
was such data that compelled Ramo and Busto (1982) to
propose that the two forms be considered the same species.
The two forms are entirely interfertile; this despite any of
the potential differences in size and soft parts discussed
above. They have long been known to interbreed in zoos
(A.P. Gray 1958), although such evidence is scarcely more
than suggestive as even distantly related animals interbreed
in captivity. However, in this case, an introduction experi-
ment demonstrated that interbreeding can occur in the wild.
Eggs from scarlet birds were introduced into Florida
(Bundy 1965, Zahl, 1967). These, and their presumed de-
scendants, interbred freely with the resident white ibises.
Birds of scarlet plumage remain in the South Florida popu-
lation, and we have been observing their interbreeding with
white birds for nearly 20 years. If they had derived from the
introductions, they would persist after several generations of
outbreeding with white birds in this way.
The next question is whether hybridization could occur in
the wild, should white and scarlet birds nest together natur-
ally. This indeed takes place in northern South America,
where Ramo and Busto (1982, 1985, 1988) and G.S. Luthin
(pers. comm. 1990) studied such pairs in the wild. They
observed no differences in nesting or courtship behaviours
in wild birds and saw considerable interbreeding between
white and scarlet forms. There is no evidence that the
offspring of these pairings would be in any way disadvan-
taged. Thus, we conclude that colour alone is not a charac-
teristic that distinguishes between two species.
Based on this basic information, two hypotheses have
recently been proposed to describe taxonomically the re-
lationship between the two forms. Ramo and Busto (1982)
proposed that these two forms be considered the same
species, but that the scarlet and the white birds be con-
sidered to represent two subspecies. Wieringen and
Brouwer (1990) proposed that the North American form
and the South American form be considered separate
species and that the South American form be considered
dimorphic.
In considering these competing proposals, it seems first
that no taxonomic characters are known that separate either
white and scarlet birds or North and South American birds
into populations that are incapable of interbreeding. This
supports the suggestion of Ramo and Busto that white and
red birds and North and South American birds are conspe-
cific.
It is not appropriate to define subspecies on the basis of
plumage colour alone, which Ramo and Busto appear to
have done, when plumages do not sort out geographically.
This supports the suggestion of Wieringen and Brouwer
that the South American form, at least, is dimorphic. It is
even possible that the North American form also is dimor-
phic, considering how regularly scarlet birds have histori-
cally been encountered in the normal range of the North
American birds. To our knowledge, consideration has never
been given to such a possibility.
The clearest concordance of the evidence with taxonomy
is to recognize a single species and two subspecies, the latter
being identified (for now) by a combination of character-

istics. The South American subspecies ruber is smaller,
usually having an all or mostly dark bill during courtship,
and males more often showing a gular sac. This subspecies
includes both white and red birds. The North American
albus is larger, usually having a part of the bill dark-coloured
during courtship, and the males seldom showing as full a
development of the gular sac as females. It should be noted
that none of these characteristics is definitively understood.
We suggest that this species is an example of the allopa-
tric divergence of the two stocks followed by secondary
contact in northwestern South America. Such a situation
would be identical to that believed to have occurred in two
other forms in the New World, the Great Blue Heron/Great
White Heron (Ardea herodias) and the Green-backed/Striated
Heron (Butorides striatus), both of which are thought to
represent subspecies in secondary contact distinguished by
plumage differences (Hancock and Kushlan 1984). The
Great Blue/Great White situation is also similar in that the
southern Florida race is dimorphic.
Our hypothesis of polytypic conspecificity should in no
way be taken to imply that the question of the relationship
between the predominantly white North American birds
and the predominantly red South American birds has been
resolved. Like all taxonomic proposals, this is a hypothesis
to be further investigated. It is our hope that detailed
quantitative studies of morphological and biochemical vari-
ation and genetic control of plumage colour will be under-
taken to determine what, if any, are the appropriate specific
and subspecific limits of populations of scarlet and white
ibises. Among problems to be addressed are documentation
of the sizes and other characteristics of birds nesting in
Central America and the birds in western South America.
Further information is also needed on the size and other
characteristics of red birds in northern populations, and of
the southern South American forms. Although such studies
would test our hypothesis of conspecificity, more import-
antly the scarlet-white ibis system could provide a mechan-
ism by which fundamental issues regarding gene flow, poly-
morphism, gene expression and speciation could be studied.
CONSERVATION
The white and scarlet ibises are well recognized birds
throughout their range, because of their impressive popu-
lation sizes and, in South America, because of their remark-
able coloration, which Beebe and Beebe (1910) called jets of
flame. The sight of flocks of white or scarlet ibises flying to
roost is one of the more memorable wildlife experiences and
may be one of the great spectacles of nature. Such roosting
flocks were prime causes of the establishment of the great
Everglades National Park in Florida in the 1930s and for the
fame of the Caroni Swamp in Trinidad. F.P. Penard and
A.P. Penard (1908) captured the scene of a scarlet ibis roost
by describing one 'as mangroves . . . splattered with blood'.
The impression made by white ibis flocks is scarcely less,
paling only in comparison with the scarlet form. These birds
are increasingly taken to be symbols of conservation activi-
ties in the areas in which they occur.
Populations remain high over most of the species' range,
American White Ibis 155
over 128000 birds nesting in the llanos alone (Ramo and
Busto 1984). However, local populations have decreased in
several areas, such as in the Florida Everglades and along
French Guiana (Spaans and de Jong 1982, pers. obs.
J.A.K.). The scarlet ibis has been placed on the endangered
species list in Brazil (Sick 1972, 1985) and little information
is available on the status of the disjunct populations in Sao
Paulo and Parana, which have reappeared and are in need
of conservation action, especially as they may be threatened
by future pollution of their newly discovered colony site
(Marcondes-Machado and Monteiro 1989, 1990).
Over its range, white and scarlet ibis adults have tra-
ditionally been hunted, and eggs and young collected for
food (F.P. Penard and A.P. Penard 1908, Bent 1926, Wet-
more and Swales 1931, ffrench and Haverschmidt 1970,
Herklots 1961, T. Barbour 1943, Ramsamujh 1990). Its
appealing taste may derive from the predominantly inverte-
brate and crustacean diet. Disturbance and killing from
hunting and other causes continues to be one of the primary
causes of reduction in numbers of breeding birds in various
areas (Haverschmidt 1968, Luthin 1984a). Hunting was
allowed in French Guiana until 1984 and hunting in
Guyana continues year round (Ramsamujh 1990, Dujardin
1990). Restrictions on hunting have been in effect in Suri-
nam since 1984 but enforcement is lax, as is the case
throughout much of the range (Held 1990). In Trinidad,
where the scarlet ibis is a national symbol, hunting has
persisted even within its sanctuaries (Hislop and James
1990). Conservation of these species in the neotropics will
require education as well as enactment and enforcement of
protective laws.
Recently a commercial threat has emerged in French
Guiana. Ben de Jong and Jean-Luc Dujardin found that an
extensive local industry had developed that produced arti-
ficial flowers from red ibis feathers (Betlem and de Jong
1983, Dujardin 1990). Despite efforts to outlaw the taking of
ibises and to find alternative sources of feathers, ibises
continue to be killed (Dujardin 1990).
With the concentration of a large portion of the species'
population in a few colonies, protection and management of
colony sites become crucial. Over the wide range of the
species, nesting sites are threatened by lumbering, wetland
reclamation, and water management. Protection is made
difficult by the inaccessibility of many colony sites, and the
species' propensity to move colony sites frequently. In
developed countries, such as in the USA, potential nesting
sites are lost through alteration of the site or nearby wet-
lands.
W. Post (1990) documented the recent extinction of the
largest White Ibis colony along the Atlantic Coast. This
colony was stable from 1974 to 1984, when over 18 000 pairs
nested, but then declined until it was abandoned by the
ibises in 1987. Causes included susceptibility to predation,
drought and floods that affected ibises more than the her-
ons. Thus complete protection and management of remain-
ing colony sites is critical in North America.
Colony site protection in Central and South America is
even more difficult because of the political and economic
situation. However, long-term sites must still be protected
156 American White Ibis
in sanctuaries and reserves. One of the most famous sites
(although no longer a colony site) is in the Caroni Swamp of
Trinidad, where many people have observed it coming into
its night roost and a tourist industry has developed sur-
rounding this nightly event.
Critical too, of course, is the protection and management
of wetland feeding habitat. Population decreases in the
Everglades are a response to alteration of the natural hydro-
logical regime of the marsh by water management practices
(Kushlan 1986a, 1987). This species has the advantage of
being able to shift easily and quickly among foraging areas
to avoid unfavourable conditions or take advantage of newly
created foraging opportunities. Wetlands will need to be
managed to maintain suitable foraging conditions for the
White Ibis and other wading birds.
The recently elucidated story of the Scarlet Ibis in the
Garoni Swamp of Trinidad is especially instructive in this
respect (ffrench 1967, 1978, 1985, ffrench and Haverschmidt
1970, Betlem 1984, Bildstein 1990a). It was not clear why
breeding ceased after this time until Bildstein, drawing
upon his studies of the White Ibis in coastal environments,
suggested that the ultimate cause was the drainage of fresh-
water marshes that served as feeding grounds during nest-
ing. This conservation story holds a message for manage-
ment of this species elsewhere, the importance of protecting
the feeding habitat. The best hope is to preserve large tracts
of coastal and inland wetland habitat.
The species serves as both an indicator of environmental
health and as a symbol of the need to undertake such
habitat conservation. Several groups are now working on
conservation plans for the species, and these must be
encouraged and implemented.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 306.
 Barefaced Ibis
Phimosus infuscatus (Lichtenstein)

Subspecies:
Phimosus infuscatus infuscatus (Lichtenstein)
Ibis infuscata Lichtenstein, 1823, Verzeichniss Doubletten Zool. Mus. Berlin, p. 75, based on
'Afeytodo', no. 365, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p.
201: Paraguay
Phimosus infuscatus berlepschi Hellmayr
Phimosus berlepschi Hellmayr, 1903, Verh. Zool.-Bot. Gessell, Wien, 53, p. 247: Orinoco River region
Phimosus infuscatus nudifrons (Spix)
Ibis nudifrons Spix, 1825, Avium Species Novae Itinere Brasiliam, 2, p. 69, pi. 86, 'Ad litora lacuum
fl. St. Francisci: Rio Sao Francisco, Bahia, Brazil

Other names:
Whispering Ibis (English); Coclito, Coclito Negro, Cocli, Ibis Negro, Zamurita, Cuervillo de Cara
Pelada, Cuercillo de Cara Afeitada, Cuervillo Pico Marfil, Cuervillo Cara Roja, Cuervo de Frente
Pelada, Curri-curri, Corocoro Negro, Tara, Corocoro Carirrojo, Yervetacho (Spanish); Tapicuru-de-
cara-pelada, Macarico-de-bico-branco, Magarico-de-cara-pelada, Macarico-preto, Tapicuru, Frango
d'agua, Chapeu-velho (Portuguese); Zwarten Clans-Ibis (Dutch)

IDENTIFICATION
The Barefaced Ibis is a small, dark ibis with green-glossed plumage and a reddish face. It is 46-56 cm
long weighing about 578 g (n = 8, range 500-600 g; Belton 1984, C.S. Luthin pers. comm. 1990).
The basic plumage colour of the adult is black but glossed with a bronzy green or blue, more so above
and on the tail and wings than below. This coloration gives the bird one of its Spanish names, Corocoro
Negro (B.T. Thomas pers. comm. 1990). The flight feathers and underwing are bluish green. The
underparts tend to a greenish brown.
The forehead, face and chin (gular area or pouch) are bare, thus providing the most common English
name, Barefaced Ibis. These and the bill are reddish, ranging from flesh to reddish brown in coloration
during the non-breeding season. The long decurved bill is slightly serrated, the same colour as the face.
The forehead may be covered with papillae or corrugations. Feathering around the face is glossy violet.
The leg and feet colours are very variable, reported as yellowish to whitish pink, reddish, grey and tan
to brown. The iris is light brown (F.P. Penard and A.P. Penard 1908), although Belton (1984) found the
iris to consist of brown, white and red concentric circles. Additional information is needed to under-
stand geographic, seasonal and developmental differences in soft-part colouring.
The three named races are described based on slight differences in plumage and soft-part colours
(Blake 1977). All have bare pink to reddish faces. The subspecies berlepschi has a relatively dark reddish
brown bill tipped in blackish and a deep red face. The nudifrons subspecies has a paler clay-coloured bill,
brick red face, and a green gloss on its hindneck feathers. Its forehead and face are rough and ridged.
The infuscatus subspecies is reported to be smaller and shorter-billed than nudifrons and to have a pale
clay- to cream-coloured bill, though Belton (1984) reported this as a greyish red. Its forehead and gular
sac are yellowish pink, and strong purple reflections gloss the feathers of the hindneck.
During the breeding season the soft parts attain their brightest colours, in some specimens appearing
bright red (Luthin 1983a). The throat area, also called the gular patch or pouch, becomes red and is
prominently displayed during courtship.

The sexes are considered to be similar, but a review of
specimens in the US National Museum suggests that males
tend to be larger then females, at least in weight and in bill
and tarsus length. With one exception, of 8 specimens exam-
ined, the bills of birds labelled Temale' were smaller than
those of 2 identified as 'male' (pers. obs. J.A.K.). Belton
(1984) recorded that a male weighed 575 g, a female 550 g,
and an unsexed (live) bird 600 g. The males also have a
thicker bill and tarsus than the females (Luthin 1983a), and
are 10% larger in bill and tarsus (C.S. Luthin pers. comm.
1990). These differences are not noticeable in the field.
Nestlings are black with a pale grey face and fleshy soft
parts. After several days, banding develops on the bill,
which, from the differing reports, may be variable. The bill
is reported to be dark with a pink tip or several pink bands
(Luthin 1983a), and to be pinkish with a dark tip (Belton
1984, B.T. Thomas pers. comm. 1990). The facial skin
above the bill is grey (or even bluish—Luthin 1983a) with a
white band, the latter being very striking within the first few
days but fading to grey by 1 week (B.T. Thomas pers.
comm. 1990, regarding berlepschi).
The bill loses its banding by fledging, although the light
patch in front of the eyes is retained. The plumage lacks the
metallic sheen of the adults, and the birds have noticeably
shorter bills (Dugand 1947).
The Barefaced Ibis is characteristically a silent bird out-
side the nesting period, having a weak, gobbling call ren-
dered 'Ghew-ghew-ghew' or £Cu-cu-cu-cu' (Sick 1985, Hilty
and Brown 1986). Hudson (1920:124) described the voice as
little sigh-like puffs of sound, hence the name he gave it, the
Whispering Ibis. As Hudson noted many years ago, the
Barefaced Ibis 159
typical quietness of this species outside the nesting season is
quite odd within the loud-voiced ibis family.
The species is more vocal during nesting. Luthin (1983a)
characterized the courtship vocalizations as falling into four
types, which he called multiple hooting, duetting, long-
hooting and pair-calling. Multiple hooting is the most com-
monly heard call, especially early on in nesting. This is a
series of several hoarse hoots, with the first sound being
louder than those following. A duet of these calls may be
performed by members of a pair, usually at the nest site or
future nest site. The duet is often preceded by a 'Whoop'
sound, the long-hoot. Pair-calling is a series of 9-30 rapid
soft hoots, by one of a pair. It is used early in nesting as a
contact call.
This dark ibis is distinctive from others in its range in its
bare, pink face and reddish bill. It can be confused with the
all-dark Glossy and Green Ibises in poor light. The Green
Ibis is larger with a dark bill, greenish gloss and short legs.
The Glossy Ibis and Whitefaced Ibis have a chestnut sheen.
The Sharptailed Ibis is larger and distinctively shaped, with
its stocky build and long tail.
DISTRIBUTION AND POPULATION
The Barefaced Ibis occurs in the tropics and parts of the
subtropics of South America east of the Andes, from Colom-
bia to Argentina, excluding the Amazon River itself. This
hiatus creates disjunct range-segments on either side of the
Amazon.
The overall range north of the Amazon is from north-
eastern Colombia, and Venezuela (eastern, central, coast-
al), Guyana, Surinam, and northwestern Brazil (Rio
Branco). South of the Amazon, the range is from central
Brazil, to Bolivia, Paraguay, Uruguay, and northern Argen-
tina south to Buenos Aires and Cordoba.
Occupying the species' range north of the Amazon, ber-
lepschi is the more northern subspecies. Nudifrons occurs in
higher lands of Brazil from central Mato Grosso, Roraima,
Maranhao, east to Piaui and Bahia, southward to Sao
Paulo. P.i. infuscatus occurs from Rio Grande do Sul (Belton
1984) to the Chaco of eastern Bolivia (Beni, Santa Cruz),
through Uruguay into northern Argentina to Tucuman,
Cordoba, Santa Fe, and northern Buenos Aires (Olrog
1963, D.A. Scott and M. Carbonell 1986).
Delimitation of the ranges of the two southern subspecies
is worthy of further clarification. They have been thought to
be contiguous, but Belton (1984) suggests that the north-
eastern littoral of Rio Grande do Sul may be a hiatus
separating the southerly infuscatus from the more northerly
nudifrons.
Populations undertake seasonal migrations (B.T. Tho-
mas pers. comm. 1990). Movements in the llanos relate to
the rainy season. Barefaced Ibises are most common there
in the breeding months and are less frequently seen in the
dry season, January to June (B.T. Thomas 1979b, E. Agui-
lera pers. comm.). They arrive in the llanos soon after the
rains begin and initiate courtship immediately. Belton
(1984) observed, similarly, that there may be some move-
ment out of Rio Grande do Sul in the cold season.
160 Barefaced Ibis
Populations can be rather substantial locally. It is one of
the most numerous species in the pantanal of Mato Grosso,
but is not found commonly in some other locations in the
region (Dugand 1947, Sick 1985). Similarly, it was the most
locally abundant ibis in the Paraguayan Chaco (Brooks
1991). Ramo and Busto (in press a) found them nesting in 7
mixed colonies in Guanare, in the Venezuelan llanos. Nest-
ing numbers per colony are low: in 1980 numbers were 22,
50, 6, 15, 100, 5 and 20 in the colonies censused. C.S. Luthin
(pers. comm. 1990) found nesting groups of similar sizes.
ECOLOGY
The Barefaced Ibis is a bird of open marshes, savanna, wet
meadows and fields. We studied its feeding habits in the
llanos, where it forages on moist ground, shallow water,
marshes and puddles, in rice fields, along streams and
ditches, and on the shore of lagoons and ponds (pers. obs.
J.A.K.). Along the coast, it feeds in tidal flats. It also uses
damp and drier sites, such as newly mowed fields and
flooded pasture (Sick 1985, Belton 1984). Although primar-
ily a bird of the lowlands, it has been observed at 2600 m
near Bogota, Colombia (Fjeldsa and Krabbe 1990).
It feeds by Walking Slowly on mud, moist land, or in
shallow water, while Probing with its bill into the soil or
shallow water. Owing to its relatively long bill, it can Probe
rather deeply in the water or soft mud but also Pecks rapidly
at the surface. We have observed them to feed very much
like the American White Ibis, by actively walking along
while Probing every 1-3 steps (pers. obs. J.A.K.). Probing
sites are typically at the edge of puddles and muddy spots.
The Barefaced Ibis typically feeds gregariously, often in
small flocks of a few birds but occasionally in flocks of up to
several hundred (Naumburg 1930). Larger single-species
flocks, as are not uncommon in Scarlet or Whitefaced Ibises,
have seldom been reported. However, one of us has ob-
served them feeding commonly among other wading birds
in rather large mixed aggregations (pers. obs. J.A.K.). In
Mato Grosso they commonly form mixed flocks with Sharp-
tailed, Glossy and Whitefaced Ibises (Naumburg 1930,
pers. obs. J.A.H.). In the llanos, we have seen them feeding
with Scarlet and Sharptailed Ibises especially, and also with
cattle (pers. obs. J.A.K.). In such cases they are brought
into interaction with other species. The Scarlet Ibis, for
example, steals prey from their Barefaced feeding associates
(Kushlan et al. 1985).
The Barefaced Ibis has been documented to eat soil
invertebrates, such as worms (probably annelids) and beetle
larvae, as well as small crabs, fish, molluscs, including
freshwater clams, and aquatic insects (Olivares 1969:80,
Belton 1984, R. Cintra pers. comm., G. Morales pers.
comm. 1990). Sick (1985) has found seeds and leaves among
stomach contents.
BREEDING
Barefaced Ibises nest in dense bushes, in colony sites that
are reused in successive seasons. They generally nest in
small groups within larger mixed-species colonies contain-
ing cormorants, spoonbills, night herons, Maguari Storks
and other ibises. They also nest in small single-species
colonies of a few to 10 nests (Belton 1984, C.S. Luthin pers.
comm. 1990). Luthin (1983a) considers the species to nest
in loose aggregates, somewhat intermediate between col-
onial and solitary nesters.
They nest in the rainy season, with some variability being
attributable to annual differences in the rainfall timing.
Typically, they begin nesting in the llanos in the rainy
season from May to late October, and in Brazil in October
and November (B.T. Thomas 1979b, Luthin 1983a, Belton
1984, E. Aguilera pers. comm.).. Nesting appears to be
prolonged and continuous through the season, as large
young can be present simultaneous with nest-building in the
same colony (Luthin 1983a, Belton 1984).
They appear to respond quickly to onset of the rainy
season, commencing nesting within 2 weeks of the beginning
of the rains (C.S. Luthin pers. comm. 1990). Courtship may
be interrupted by windy conditions.
Pair-formation was studied by Luthin (1983a) in the
llanos, and most information on the process is derived from
this important study. The males gather in display groups in
variable locations with the composition of the display party
changing frequently. Fighting among males seems to be
minimal. Erect Postures and Display Shakes occur as other
males approach too closely. There are occasional Sparring
and Supplanting displays, but these are not common.
The birds display more often in the mornings and late
afternoons, departing the display grounds in midday except
during the peak of the courtship period. Birds arriving at the
display area, land with a characteristic 'whoosh' of their
wings. Luthin reported seeing both sexes displaying.
The most common behaviour is Display Preening,
directed to all parts of the body, most often to the back with
the wings held open and one of them drooped. The feathers
on the neck and back may be raised a bit, and the red skin of
the gular patch is often flared. During this display, the bird
may also perform a Head Rub, touching, holding or rubbing
the top of its head to its back. Both members of a pair may
Display Preen together, or one may lead and the other
follow. Luthin reported that Display Preening by the male is
more vigorous and exaggerated, and the female seldom
droops her wings. Allopreening between the members of the
pair appears to be rare.
The female will use a Stick Shaking display when she
lands next to the displaying male. This is followed by a Stick
Shake by the male. It appears that this is primarily an
appeasement display.
A pair may engage in duetting, which becomes more
synchronized as the pair-bond develops. This is often fol-
lowed by Stick Shaking. Pairing birds often stand erect, side
by side, with the males standing taller.
An Erect Posture is assumed in response to disturbance.
Display and nest-sites are defended with a Sparring display,
accompanied by a high intensity multiple-hoot and exten-
sion of the gular area. At this stage, fights may occur
between displaying males. To displace another bird, an
attacking male will use Supplanting display, in which it flies
towards or drops down on the other bird with feet and wings

extended. A male encroached upon by another may raise
and shake the feathers of his back and neck. Luthin (1983a)
called this a 'Ruffle display' but it may be considered a
variation of the Display Shake.
The nest-site is selected after initial pair-formation. The
new pair moves from the highly visible display perch to a
more suitable nest-site within the canopy, where pair-
formation displays and site defence are continued. At this
point, defence may be quite aggressive, including Sparring
displays, with the body tipped towards the foe, wings spread
and feathers raised. In this posture, a bird will walk towards
the intruder and may even chase it away, following it from
his nest site. Females give a variation of the Sparring display
directed towards intruding birds, while remaining in sitting
position on the nest. This posture, which shows the red chin
patch, is held for as long as the intruder is present.
Luthin observed that this intense territorial behaviour
keeps closely synchronized birds from nesting near each
other. After nesting gets underway, a pair allows establish-
ment of new nests nearby, so neighbouring nests are usually
in different stages of nesting.
Early copulation takes place at the display site, later
copulations at the nest site. Copulation is preceded by Stick
Shaking and Bill Shaking, which in this species is more
appropriately called Bill Swinging. The male comes up to
the female, and may put his head over hers, Stick Shakes,
and then begins to slowly swing his head back and forth
bumping the female's bill. Upon placing a foot on the
female's back, the male continues this Bill Shaking (Swing-
ing) by taking the female's bill in his. During copulation
both sexes raise their feathers and flare their gular sacs.
Copulation may be followed by tail wagging or ruffling the
body feathers.
Promiscuous matings are common, perhaps a reason why
nesting birds so strongly resist the establishment of neigh-
bouring nests until after incubation begins. C.S. Luthin
(pers. comm. 1990) found one such event occurring when
young were in the nest. These matings appear to be forced.
The male gathers material for the nest, which is built by
the female. Early twigs are used for Stick Shaking rather
than nest-construction. Later, both birds simultaneously
Stick Shake as the twig is handed from the male to the
female. Nest-construction may continue into incubation.
The nest is placed rather inconspicuously deeply within
the branches of thick, often thorny, shrubs and trees usually
over water (C.S. Luthin 1983a, pers. comm. 1990). The nest
is a rather small, flat platform of sticks, 30-35 cm wide
(Meyer de Schauensee and Phelps 1978:23).
Eggs are blue-green and unmarked (B.T. Thomas pers.
comm. 1990). Nests with 2-6 eggs have been observed in the
llanos (Luthin 1983, Ramo and Busto in press), 3-4 being
most common; the average is 3.8/nest in the Venezuelan
llanos (Luthin 1983a). These eggs averaged 47.3 X 34.5 mm
Barefaced Ibis 161
in size. Luthin (1983a) suspects that egg dumping may
occur, as suggested by the appearance of additional eggs
rather late in incubation.
Both parents incubate, over a period of 21-23 days, and
attend the nest until the young are about 1 week old. In
addition to the threat of intruding ibises, eggs are at risk to
predators. For example, eggs are eaten by the boa (Constric-
tor constrictor), the tegu lizard and predatory birds (B.T.
Thomas pers. comm. 1990, Luthin 1983a). Luthin observed
that adults did not interfere with the lizard eating their eggs.
The threat of predation and lack of nest defence undoubt-
edly accounts for the secluded positioning of the nest.
Exchanges at the nest occur up to 6 times per day and are
accompanied by a Greeting display in which the incubating
bird gives a long-hoot while the bill is pointed upward and
the head back, followed by Stick Shaking. When the reliev-
ing bird arrives, both duet and display together. Males may
also continue to supply twigs. Later in nesting, exchanges
include less display.
Young hatch asynchronously and are altricial. After sev-
eral days all chicks appear to be the same size, so brood
reduction may not be common. Young are silent, except for
the light buzzing call given at feedings by older nestlings
(C.S. Luthin pers. comm. 1990). Like the eggs, nestling are
at risk to predation, and the young will leave their nest in
response to disturbance. They depart the nest permanently
in 22-27 days and fledge at 27-30 days (Luthin 1983a).
TAXONOMY
Based on his observations of behaviour, C.S. Luthin (pers.
comm. 1990) concluded that the Barefaced Ibis appears to
have no close relatives. Resolution of the question of its
relationships to other ibises will require additional study.
Similarly, additional study is needed of the named subspe-
cies. The highly variable soft-part colours, especially, need
to be more carefully evaluated, as do the range and degree of
differentiation between the two southern subspecies.
CONSERVATION
The Barefaced Ibis is widespread, locally quite common
and under no specific threat. However, a lack of detailed
information on its habitat needs, food, and migratory and
dispersal strategies would hamper the development of con-
servation plans, should these become needed locally.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 307.
 Glossy Ibis
Plegadis falcinellus (Linnaeus)

Tantalus Falcinellus Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 241; based on 'Numenius rostro arcuarto'
of W.H. Kramer, 1756, Elenchus Veg. Animal. Austriam Inferiorem Observatorum, p. 350, and 'Le
Gourly verd' of Brisson, 1760, Ornith., 5, p. 326, fig. 2 (immature): Austria Italia ( = Italy)

Other names:

Black Curlew (English); Corocoro Gastano, Coco, Coco Prieto, Coco Oscuro, Ciguena, Morito,
Coclito Moro, Coclito Moreno (Spanish); Ibis Falcinelle, Pecheur, Ibis Noir, Pecheur (French);
Sichler, Brauner Sichler (German); Zwarte Ibis (Dutch); Mignattaio (Italian); Tambala Dahakatti,
Kokka (Sinhalese); Karuppu-Kottan (Tamil); Voromainty, Renikato (Malagasy); Karavaika
(Russian)

INTRODUCTION
The widespread Glossy Ibis is a small dark ibis, standing 56-63 cm tall and weighing 500-800 g. It is
characterized by having a striking bronze green tinge to its otherwise dark plumage.
During the non-breeding season, the adult's head and neck are sooty brown with thin white streaking
originating from white-margined feathers. A shiny chestnut gloss is particularly obvious on the feathers
of the back, wings, tail and front of the head. The flight feathers are greenish black. The exposed skin of
the face is blackish purple, the iris is dark grey to brown, and the bill dark brown to blackish. The legs
are dark brown.
The sexes are similar in plumage coloration, but the male is larger in all body measurements. The
male's total length averages about 63.5 cm, contrasted with 55.3 cm for females (Dementiev and
Gladkov 1951). The differences are so noticeable that they can be used as field marks.
A partial pre-nuptial moult occurs, and, during the courtship season, the plumage increases in
radiance. The sheen becomes a rich chestnut on the neck, mantle, shoulders and underparts, and purple
and green on the head, upperparts, tail and wings. The lores remain dark brown, but the face turns pale
blue at courtship, fading to dark purple thereafter. A distinctive white to pale blue line develops on the
skin bordering the feathering of the forehead and side of the face but does not extend behind the eye or
under the chin. A complete post-nuptial moult occurs prior to winter.
The nestlings are sooty black. They have a red patch of skin on the crown, which fades over the first
week, as well as an accompanying patch of white feathers on the head and throat, the white differing in
extent among individuals. The bill is pink with a variable amount of black banding. The lores, edge of
mouth, legs and feet are pink; the iris is greyish brown.
The juvenile plumage is dark dusky green, but not glossed to the extent of the adult and without
chestnut overtones. The head and neck are greyish brown, mottled with white on the face and streaked
with white on head and neck; the small white patches above, on the crown, fray off rapidly. The
undersides are brownish, with a faint greenish bronze tinge. The bill, initially pink, gradually darkens,
starting at the tip. Legs and feet become a dark olive-grey, the iris a darker brown. With moult, the
plumage becomes more like the adult's, acquiring a full sheen over the first 2 years.
The Glossy Ibis has a call similar to the American White Ibis, a sheep-like 'Huu-huu-huu-huu'. An
adult uses a guttural 'Graa-graa-graa' when approaching the nest and engages in mutual cooing sounds
during courtship.
The flight is rapid wing-beats interspersed with gliding in small tight flocks or in elongated V flights.
Birds often fly long distances to and from feeding grounds. They may be seen in acrobatic manoeuvres,
164 Glossy Ibis
rising high and then falling out of the air with their legs
dangling.
The Glossy Ibis looks black from a distance and can be
confused with similar dark birds under such conditions. It
might at a distance be confused with a curlew, but the
Glossy Ibis is darker, larger, carries itself more upright, and
has longer appendages.
Adults are not safely distinguished from the Whitefaced
Ibis at a distance in the field, even during the nesting season
when both birds have white bands on their forehead. How-
ever, the Glossy Ibis is larger; its facial skin is bluish white
rather than reddish; its legs are dark rather than reddish;
and the white, facial line is the result of skin pigmentation
rather than feathering as in the Whitefaced Ibis, nor does it
extend behind the eye as in the latter species. Juveniles and
wintering birds of the two species may not be distinguish-
able in the wild at all, although juvenile Glossy Ibis have
darker upperparts than juvenile Whitefaced Ibis.
The Glossy Ibis might be confused in the field with
several dark South American ibises. However, it is larger
and slimmer than the crested Green Ibis and has darker,
longer legs than the Barefaced Ibis. Juvenile American
White Ibises have lighter undersides. In many cases, close
study is required to distinguish these dark ibises, which may
feed together. In Asia, the Black and Giant Ibises are
bigger, and the former differs in having white wing patches.
In Africa, the species can be distinguished from the
Hadada by the latter's duller plumage, jerky flight and
distinctive call. It can be told from the other Bostrychia ibises
by its lack of a crest and by its glossy sheen. The dark
Waldrapp Ibis has a bald head and a neck ruff.
DISTRIBUTION AND POPULATION
The Glossy Ibis is the most widespread species of stork or
ibis, occurring in parts of Eurasia, Africa, Australia, and
North, Central and South America, and on various islands
(Keve 1967).
In the New World, the breeding range encompasses east-
ern North America, from New Brunswick, Canada, and
Maine through Florida and Louisiana, sporadically inland
as far as Arkansas, USA. It has also been recorded as
nesting in the Greater Antilles (Cuba, Hispaniola, Puerto
Rico), Central America (Palo Verde in Costa Rica), and
northern South America in Venezuela (Aragua, Zulia,
Apure) (Meyer de Schauensee and Phelps 1978).
In Eurasia, it breeds particularly in southeastern Europe
through Russia, in northeastern Italy, Austria, Yugoslavia
(Sasko Lake), Albania, Greece, Bulgaria, Romania, Molda-
via, the Ukraine, southern Russia (Kuban and Terek
Rivers), Armenia, Turkamen, Kazakh, Turkey, Israel,
Iraq, Iran, Pakistan (Sind), India (Punjab, Uttah Predesh,
Orissa, Madhya Predesh, Manipur, Assam), Bangladesh,
Burma, Vietnam (Minh Hai), Indonesia (West Java, Kali-
mantan, Irian Jaya, probably Sulawesi), the Philippines
(Mindanao, Luzon), and northern and eastern Australia.
In Africa, it nests locally in Tunisia, Morocco, Mali,
Kenya, Tanzania, Zambia, Zimbabwe, Botswana, South
Africa, Namibia, Angola and Madagascar.
The species is highly nomadic and shifts its nesting sites
quickly. As a result, it has been documented as breeding in
only a small portion of its overall range, sometimes for only
a few years in any specific location. It also nests in relatively

inaccessible areas such that ephemeral colony sites can be
overlooked easily. In any one year, it undoubtedly nests in
more places and over a wider area than is documented.
The nesting range of the Glossy Ibis has changed mark-
edly in the present century. In North America, the breeding
range has greatly expanded. Although known from 1817, it
was first found breeding on that continent in the late 1800s.
The known nesting population in Florida grew to 1200 pairs
over the next quarter-century, after which it declined while
simultaneously expanding its range elsewhere in the USA: it
nested in Louisiana in the 1930s, in Virginia and New
Jersey by the mid 1950s, New York and Arkansas by the
1960s, Maine in 1972, and New Brunswick, Canada, in 1986
(Semple 1932, Sprunt 1933, Bock and Terborgh 1957, Hail-
man 1959, P.W. Post 1962, Hanebrink and Cochran 1966,
Finch 1973, McAlpine et al. 1989). Populations apparently
began expanding and increasing in northern South America
by the mid 1960s (Gochfeld 1973). Expansion also may have
occurred in Africa and the Middle East, with nesting first
reported in South Africa in mid 1950s and Israel in 1969
(Paz 1987). It has recently been reported to nest in Pulua
Dua, West Java (Milton and Marhardi 1985).
In other areas, the range has contracted markedly, espe-
cially in western Europe and North Africa. It no longer
nests in southern Spain, southern France, Sicily, Austria,
Morocco, northern Algeria or Egypt (Cramp 1977, Pineau
and Giraud-Audine 1977). Previously it was also reported to
nest in Sri Lanka and northwestern Mongolia.
The species disperses widely outside its more usual range,
occurring as a straggler in many additional places. It has, in
fact, been seen in most temperate to tropical areas at some
time. After the breeding season, it wanders as far afield as
Iceland, Bermuda, Azores, Madeira, Cape Verde Is., Sao
Tome, Faroe Is., Great Britain, Scandinavia, Germany,
Poland east through Tibet, China (Chekiang, Fukien,
Kwangton including Hainan), Sri Lanka, Hong Kong,
Cambodia, Borneo, Moluccas, New Guinea, New Zealand
and Tasmania. In the New World, it wanders to Quebec,
Newfoundland, Nova Scotia, the Bahamas (Great Inagua,
Cay Lobos), the Cayman Is., Puerto Rico, Jamaica, the
Virgin Is., Lesser Antilles (Guadeloupe), eastern Mexico,
through Central America (Honduras, Costa Rica, Panama)
and northern South America (coastal Colombia and Vene-
zuela).
Dispersive and nomadic wanderings turn into a true
migration in autumn. In North America, Glossy Ibises from
the eastern USA winter in Florida and the Gulf Coast and
also move through Florida to the West Indies (Bahamas,
Greater Antilles, northern Lesser Antilles), Central Amer-
ica and northern Venezuela. Eurasian birds winter along
the Mediterranean in southern Europe and the Middle East
through western USSR to Iraq, and Iran through to India,
and also in northern Africa from Morocco through the Nile
Valley and the Sudan. They also cross the Sahara and
follow the Nile valley into central and east Africa. Birds
from eastern Europe move either to East Africa or to India.
In Asia, ibises winter through India to Southeast Asia, the
East Indies, and Australia. It is likely Australian birds
winter in large numbers in Irian Jay a (M J. Silvius and
Glossy Ibis 165
W.J.M. Verheugt pers. comm. 1990). There may be mi-
gratory movements in Africa as well (Tarboton 1977a). The
return north is rather protracted in the spring and many
begin to nest after the other birds in mixed colonies.
The Glossy Ibis is never abundant though widespread, as
individual populations are small and changeable, and in
most areas is considered to be locally rare. In Australia,
Glossy Ibises wander nomadically and do not use regular
nesting sites (K.W. Lowe 1983). Colony sites and groups of
colonies tend to number in the hundreds to low thousands of
nesting pairs (Luthin 1984a). However, over its entire range
the population is considerable.
Available information reflects changing local abun-
dances. In Greece, the population has decreased from 1840
breeding pairs before 1970 to 50-71 pairs in 1985-86 (Cri-
velli et al. 1988). Colonies of over 12 000 birds were found in
the Volga Delta in the 1930s, but few remain. Flocks of over
1000 birds are still seen in the USSR.
Our observation in northern India suggests that,
throughout the Asian region, the Glossy Ibis is now very
sparsely distributed, and is absent from many apparently
suitable areas, probably due to the lack of well-protected
breeding sites. It occurs on Sulawesi in fairly large numbers
and in Irian Jaya in flocks of up to several thousand
(C.M.N. White and M.D. Bruce 1986, Uttley 1987, MJ.
Silvius and W.J.M. Verheugt pers. comm. 1990).
Even where it occurs more abundantly, year-to-year vari-
ation in breeding numbers may be substantial. Comprehen-
sive censuses along the North American east coast found
12300 breeding Glossy Ibis in 1975 (Custer et al. 1980).
The following year, 25% fewer birds were found, demon-
strating the rather typical variability in nesting effort and
distribution from one year to the next.
The present status of the species and nesting populations
are unknown in many areas, especially in China, the Philip-
pines, Madagascar, the USSR, West Africa and northern
South America. More information is needed in these areas.
ECOLOGY
Glossy Ibises are marsh birds, using fresh and saltwater
marshes throughout their range. They particularly favour
lake and river-edge marshes, such as are found in river
deltas, and their breeding distribution often follows water-
courses. Along the coast, they frequent saltmarshes and
estuaries. They are particularly found in irrigated farm
fields, especially rice paddies, throughout the range, and
they are also attracted to muddy substrate exposed by
falling water levels in wetlands and reservoirs.
Population shifts have been in response to drainage of
river flood-plain, which destroys nesting and feeding habi-
tat, and to development of reservoirs, which provides new
nesting sites. The core of the Old World populations is in
the USSR, where the species occupies the deltas of rivers
emptying into the Black, Azov, Caspian and Aral seas, and
islands in reservoirs in that area.
The Glossy Ibis feeds in very shallow water, generally in
small flocks, often well dispersed. It feeds by Walking
Slowly while Probing rapidly into the water and soil, usually
166 Glossy Ibis
in mud away from plant cover or in short vegetation. Few
other observations of foraging behaviour have been made.
The Glossy Ibis often feeds in mixed-species aggregations
of wading birds. When feeding in mixed-species flocks, it is
often followed by commensals such as egrets, which appear
to benefit by catching prey items frightened into flight by
the probing ibis flock (Erwin 1983). In Florida, we have
seen them feeding alone in mixed flocks, especially with
Snowy Egrets (Egretta thula) and American White Ibises.
However, we have also seen them feed alone, especially
males. In northern Australia, they feed with Royal Spoon-
bills and on the fringes of areas where Blacknecked Storks
feed. In the Bharatpur Reserve in northern India, they are
found only in small numbers and feed solitarily in the
shallow marsh edges of pools.
The food is primarily invertebrate, including insects and
insect larvae (dragonflies, crickets, water beetles, scarabs,
weevils, soldierflies, leafminer beetles and flies), worms,
snails, spiders, and crayfish and other crustaceans. Less
frequently, fishes, frogs, tadpoles and snakes are consumed
in numbers, when these become available. The diet varies
seasonally, depending upon what is accessible. Tadpoles are
eaten when abundant, and also locusts when swarming.
BREEDING
The Glossy Ibis is primarily a marsh bird, usually nesting in
dense reeds near its preferred feeding areas. Secluded lakes,
thickets of reeds, river oxbows and islands of vegetation are
favoured nesting sites. Reed nesting is probably the primi-
tive nesting habit, but the species also uses a wide range of
nest sites including trees, bushes or the ground.
Nesting is usually in mixed species colonies with other
waterbirds, but Glossy Ibises have also been reported to
nest singly (Le Souef 1917). In Florida, we have found them
nesting intermingled with the American White Ibis and
Cattle Egrets (Bubulcus ibis). In Kenya, we found their nests
to be scattered throughout a mixed colony consisting mainly
of egrets. In Australia, 1000 Glossy Ibis have nested with
40000 Strawnecked Ibises, 1000 Oriental White Ibises,
Yellowbilled Spoonbills, Royal Spoonbills and other wading
birds (K.W. Lowe 1983). They also nest with cormorants
and pelicans.
The nesting season is generally in the spring, May to
August in eastern Europe and eastern North America, and
earlier further south. In southern Florida, we have found
them nesting as early as March, but even here they are more
usually associated with summer nesting colonies. Such vari-
ability in nesting timing is not unusual. In Africa, on the
Tana River, Kenya, we counted 50 pairs nesting in the
winter rains and at the Garsen Colony Site, we found 60
pairs nesting in June of the same year.
Courtship behaviour is virtually undescribed (Baynard
1913a,b). After pairing, a Greeting display continues to be
given throughout nesting. It involves considerable mutual
bowing, rubbing of bills together, and guttural cooing vocal-
izations. Prior to nest-relief, the incoming bird gives a call
some distance from the nest and is answered by the mate.
Later in nesting the incubating bird may leave at the call,
just prior to the arrival of the relieving bird without inter-
action.
The nest is built by both birds and both collect material.
However the male brings most of the nesting material. Ibis-
built nests are relatively bulky for their size, averaging 30-
40 cm wide, and 4-27 cm thick. They are made of reeds or
twigs and lined with green leaves or grass, which are con-
tinually added throughout nesting.
Within a colony site, the birds choose to nest in the denser
available vegetation and to be rather dispersed throughout
the colony, as noted above. Most typically, they place their
nests low to the ground, especially on marsh vegetation or
even on the ground. In Kenya, we found the nests built in
heavy-foliaged trees and tall bushes, near and beneath the
nests of African Spoonbills and African Openbill Storks. In
Australia, they have been reported nesting on the trunks
and branches of knocked-down lignum trees (K.W. Lowe
1983). They also reuse nests built by egrets, and these are
positioned higher than the nests that the ibis build for
themselves (Burger and Miller 1977).
The eggs are deep, solid blue, fading to a lighter blue
during incubation. The clutch size is 2-6, averaging about
3, and egg dumping may occur (Jozefik 1960b). Eggs aver-
age 53 X 36mm in size and weight 31-39 g (K.W. Lowe
1983). Egg-laying within a colony may be spread over 5 or
more weeks (K.W. Lowe 1983), and there is evidence of
renesting within a breeding season.
Incubation, by both sexes, takes 21 days, the larger male
taking the daytime shift. Chicks hatch asynchronously, and
a substantial brood reduction occurs, with an average of 1 or
fewer chicks fledging per nest in most studies, the reduction
being attributed to starvation and predation (L.M. Miller
and J. Burger 1978, K.W. Lowe 1983).
Both adults care for young, one parent being present
during the first week. We found nest-relief to be done furti-
vely, with the relieving bird creeping along branches well
hidden until the change-over occurred. The young are fed
by regurgitation and are not as aggressive as American
White Ibis chicks. Young beg by touching the parent's bill,
which inspires regurgitation: as the chick grows, it will place
its entire head into the parent's gullet.
Chicks are able to move from the nest in 2 weeks to form
creches in nearby branches or on the ground, but continue
to be fed at the nest site. They are surprisingly docile and
are with difficulty described in any other way than
Affectionate'. They fledge in 25-28 days. Depending on the
colony, they may stay in the colony for another 3 weeks or
fly to the feeding grounds with the parents soon after fledg-
ing. Young fledge at less than adult size, which is not
attained for 6 months.
TAXONOMY
The Glossy Ibis is closely related to the Whitefaced Ibis.
The two species can interbreed in captivity and are some-
times considered to be conspecific (Parkes 1955, Palmer
1962). However, the two forms nest sympatrically in Loui-
siana and Alabama, USA (Duncan and Johnson 1977), and
so appear to be specifically distinctive. The unusual situ-

ation of having such closely related forms nesting together in
North America is best understood by noting the lack of
subspecific differentiation between Old World and New
World populations. We believe that the Glossy Ibis is a
natural invader of North America, perhaps within the his-
toric period.
The Glossy Ibis is often divided into two subspecies with
P. f. peregrinus being the form found in Australia, Indonesia,
and Madagascar. It was described as having smaller wings
and tarsus, but this distinction has not held up to closer
scrutiny (Amadon and Woolfenden 1952, Vaurie 1965).
Although we can reject this subspecies, the apparent uni-
formity of the Glossy Ibis over such a wide range deserves
careful study.
CONSERVATION
In the Old World, nesting and feeding sites are threatened
by drainage and development of wetland habitats, and
nesting sites are enhanced by reservoir development (Olei-
Glossy Ibis 167
nikov et al. 1975). The situation in the southern USSR is
typical. Populations there decreased due to development
and then stabilized due to the creation of reservoirs and
protection breeding sites (Oleinikov et al. 1975). In the
Volga Delta, the high populations of 1930s have more re-
cently declined to a single pair by the 1960s (Cramp 1977).
The actual status of this species is very poorly known
owing to its tendency to shift nesting sites so frequently and
quickly. Apparent population decreases in one area are
sometimes found to correspond to increases in other areas.
Perhaps the most important contribution to the conser-
vation of this species is to conduct additional studies of its
biology, movements and population structure, and to pro-
tect the reedmarshes on which it depends.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 308.
 Whitefaced Ibis
Plegadis chihi (Vieillot)

Numenius chihi Vieillot, 1817, Nouv. Diet. Hist. Nat., nouv. ed., 8, p. 303; based on 'Cuello jaspeado',
no. 364, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p. 197: Paraguay
and campos of Buenos Aires, Argentina

Other names:

Whitefaced Glossy Ibis, Black Curlew, Bronze Ibis, Black Curlew (English); Atotola, Morito
Cariblanco Atotola, Acolote, Cuervo, Gallareta, Cuervo de Pantano, Cuervo de Canada, Cuervillo de
Canada, Cuervo de Pantano Comun (Spanish); Carauna, Tapicuru, Magarico-preto (Portuguese)

INTRODUCTION
The Whitefaced Ibis is a small (46-56 cm, 450-525 g), dark ibis, with glossy purple, green and bronze
reflections, and, in the breeding season, white feathers around the bill. This is a striking bird, which,
because of the brilliance of its metallic sheen, is often called 'a work of art'.
Outside the breeding season, the adult's base colour is a dark maroon-brown. The underparts are
deep brown with vinaceous overcast and minor dull dark green reflections on the breast (Belknap 1957).
The head and throat are streaked in white. The legs and face are reddish, and the iris reddish brown.
The sexes are similar in plumage but the male is larger, heavier and has a longer bill. Bill length can
be used to tell the sexes apart in the field, as overlap in size between them is minimal (Belknap 1957).
There is considerable variation among individuals, however, and the South American forms may be
smaller than their North American counterparts.
The full plumage brilliance is achieved in the nesting season following a pre-nuptial moult. This
occurs in April and May in North America and August and September in South America (Belknap
1957, S. Narosky pers. comm. 1990). The head, neck, upper back, wing-coverts and undersides assume
a deep, dark, chestnut-maroon coloration. The dark uppersides reflect iridescent purple, violet and
green. The dark flight feathers are metallic green and bronze, whereas the wings have purplish
overtones.
The bare skin of the face becomes reddish or purplish. A line of white feathers, variable among
individuals, separates the forehead and the face, extending completely around the eye. The chin is also
white, and this colour sometimes extends down the throat. The iris is reddish brown. The bill, legs and
toes are blackish grey. Complete post-nuptial moult occurs in late summer, July and August in North
America.
Plumages and coloration of young have been well documented (Belknap 1957, S. Narosky pers.
comm. 1990). Hatchlings are covered with a sparse brownish black down, darker and denser on the
uppersides than elsewhere (Belknap 1957). The legs and bill are pink orange. The short, straight bill
has a single central band, which, like the tip, is lead coloured. The bare crown of the head is pink with
cinnamon and white down. The literature suggests to us that there may be more white on the crest of
North American nestlings than on South American birds. The legs are black with bluish grey nails
having pinkish tips. The iris is a very dark greyish brown.
The juvenile plumage begins to develop within 2 weeks. It is basically a shiny dark green, initially
retaining traces of down, especially on the head. By the third week, the bare area of the top of the head
develops a light orange colour. The head and neck acquire white mottling, and so are lighter than in the
adult. The body colour lacks the rich chestnut hue. Upperparts are glossed with greenish highlights;
170 Whitefaced Ibis
underparts are lighter, with faint purplish or greenish
sheen, depending on the light. The iris is brown, and the
legs and bill are black. The bill is a whitish pink, with black
rings at the base, middle and tip. Newly fledged young are
smaller than adults and have noticeably smaller and less
downcurved bills. The bill stripes may be retained into the
first winter, during which the bill darkens. The post-juvenile
moult begins in late summer and fall, the birds developing a
brownish plumage with purple-chestnut sheen and the
streaked brown and white neck of the wintering adult.
The Whitefaced Ibis flies strongly, quickly and gracefully,
alternating bouts of quick flapping with gliding. It flies with
its head and neck completely extended, appearing black
from a distance. However, at closer range or when the sun
hits the birds, the bronze and green sheen become obvious.
They fly in small to large flocks, often in ever changing' V
or linear formations. Flocks can number in the hundreds,
and dusk roosting flights can be spectacular. These birds
also soar, especially on migration, and can engage in specta-
cular aerial manoeuvres, in which they descend rapidly in a
twisting path, falling nearly to the ground, before rising
again.
When in flight or on the ground, the usual call is a loud
nasal (Ka-onk, ka-onk, ka-onk', which Hudson (1920) lik-
ened to human laughter. In courtship, both sexes give
repeated 'Geek' and other calls.
In the wild, it is difficult or impossible to distinguish
between juveniles and some wintering individuals of the
Glossy Ibis and Whitefaced Ibis. In the breeding season, the
two species are distinguished by the difference in plumage
sheen, bare-skin colour, and the character of the eye-line.
However the latter is not a reliable feature under most field
conditions, because the Glossy Ibis has a similar white line
of skin during that time of year. The white feathering of the
Whitefaced Ibis does extend all the way round the eye,
which the white skin patch of the Glossy Ibis does not. The
otherwise dark juvenile American White Ibis has white
undersides.
The glossy sheen and facial coloration will also dis-
tinguish this ibis from other dark ibises in its South Ameri-
can range. The Whitefaced Ibis is often seen, and can be
confused, with the Barefaced Ibis, which is dark, less glossy,
and has a pale face and bill (S. Narosky pers. comm. 1990).
In flight the two are easily told apart, as the legs of the
Whitefaced Ibis extend well beyond the tail whereas those of
the Barefaced Ibis cannot be seen. The Puna Ibis is dis-
tinguished by its black undersides (Sick 1985).
DISTRIBUTION AND POPULATION
The Whitefaced Ibis has an exceptional breeding distri-
bution with two entirely disjunct nesting populations, one
centred in the Great Basin of western North America and
the other in the pampas of central and southern South
America. The reasons for this odd distribution are unclear
(R.A.Ryder 1967).
The northern population is concentrated in the Great
Basin states of the United States and Canada, including the
eastern Oregon, southern parts of Idaho, Wyoming, Mon-
tana, North Dakota, Alberta and Minnesota, east to Ala-
bama, west to central California (San Joaquin Valley), and
south to New Mexico, Texas, Louisiana, and Colima, Zaca-
tecas, Veracruz and Tabasco, Mexico (R.A. Ryder 1967,
Steffen et al. 1978, Skaar 1975, Salt and Salt 1976, Loke-
moen 1979, Portnoy 1981, Ivey and Severson 1984, Findholt
and Berner 1988, Stahlecker 1989, D.M. Taylor et al. 1989).
The southern population nests in eastern Bolivia, Para-
guay (Tebicuari), Uruguay, Brazil (Mato Grosso, Sao
Paulo, Rio de Janeiro, Santa Catarina, Rio Grande do Sul),
central and southern Chile, and south to Argentina (Los
Yngleses, Ajo, Buenos Aires, Rio Negro). A disjunct popu-
lation occurs in coastal Peru. Even within this larger range,
breeding populations are quite localized. The Whitefaced
Ibis is, in fact, transient over a large part of its overall
breeding range. The small Chilian population has appar-
ently not nested in the last decades (Schlatter pers. comm.
1990).
The northern population is migratory. They disperse
after nesting to various feeding locations, some of which
they remain at until fall (Kotter 1970, D.M. Taylor et al.
1989). Wintering birds also are found widely from southern
California, Colorado, Arizona, coastal Louisiana, Texas,
Baja California and Mexico. Most of the wintering popu-
lation appears to be focused in Mexico, including Jalisco,
Michoacan, Nayarit, Guanajuato, Queretaro and the
Federal District. Wintering birds also have been reported
from further south in Central America, in Guatemala, El

Salvador and Honduras, but considering the difficulty of
identifying the species, additional information is needed on
this point (Monroe 1968:58).
According to S. Narosky (pers. comm. 1990), the
southern population in Argentina does not migrate; this
conclusion is contrary to that of Hudson (1920). Juvenile
birds in South America do disperse after nesting, however.
Banded young have been recovered as far as 1800 km from
their natal colony, and have moved as far as 1400 km within
a few months (Olrog 1975, Sick 1985).
Individuals also wander, especially prior to nesting in
response to dry conditions. Birds have been found as far
north as southern British Columbia, Alberta, Saskatchewan
and Manitoba, as far east as Ohio, New York and Mary-
land, as far south as Florida and Costa Rica, and as far west
as Hawaii. In South America, stragglers are reported as
high as 4300 m in the mountains of Bolivia (Fjeldsa and
Krabbe 1990) and as far south as the Straits of Magellan.
We consider that the previously accepted records of
Whitefaced Ibis nesting in Venezuela and Colombia (e.g.
American Ornithologists' Union 1983) refer in fact to the
Glossy Ibis (Gochfeld 1973). The single nesting record from
Florida appears to be valid (Brewster 1886, Parkes 1955),
but we conclude that it was an isolated incident. These
locations are therefore not included as part of overall range
of the species.
The Whitefaced Ibis is a common species and can be
abundant locally. However, nesting colony sites are widely
dispersed within its overall range. In the mid 1960s, North
American nesting populations decreased precipitously due
to pesticide contamination, especially from feeding in agri-
cultural fields. At that time, R.A. Ryder (1967) estimated
10000 breeding pairs remained in the USA. The Texas
nesting population decreased from 9200 pairs to 2100 pairs,
from 1969 to 1974 (K.A. King et al. 1980).
However, populations remained high in most of the
species' southern range. They are abundant on the pampas
of Argentina (Hudson 1920); Weller (1967) reported 15000
nests from San Antonio, Argentina, and Belton (1984)
reported 12000 nests in Rio Grande do Sul, Brazil. In the
marshes of Rio Saladillo, Cordoba, Argentina, 20 000 win-
tering Whitefaced Ibis were reported (D.A. Scott and M.
Carbonell 1986:20).
This species has recently been increasing in North Amer-
ica, reclaiming and surpassing its historic breeding range,
north to Oregon and Canada and east to Alabama. By the
mid 1970s over 6000 pairs of nesting Plegadis ibises (almost
certainly this species) were counted in Louisiana, and
another 5300 pairs occurred in Texas (Portnoy 1981, K.A.
King et al. 1980). In 1985, 5000 breeding pairs were studied
at a single colony in Nevada (Bray and Klebenow 1988).
They are known to nest as far south as the Tampico lagoons
(100 pairs) and the Tabasco Lagoons (500 birds counted) in
Mexico (D.A. Scott and M. Carbonell 1986).
ECOLOGY
A marshland bird, the Whitefaced Ibis is found in fresh and
saltwater marshes and wet grassy areas, along ponds, rivers
Whitefaced Ibis 171
and reservoirs, and in open grassland. They feed in water
less than 20 cm deep, in damp meadows, and on open dry
grasslands. In recent years, they have accommodated to
flooded pastures and irrigated fields, especially rice fields.
In the Great Basin, they feed often in agricultural fields,
mostly in puddled water in recently tilled and irrigated
alfalfa fields (Bray and Klebenow 1988). They can travel
considerable distances to feeding areas, over 30 km from
colony sites in Argentina (Weller 1967). They are, however,
quite dependent on rainy conditions to provide foraging
habitat, and nesting success depends on weather, failing in
drought years and succeeding in wet years (R.A. Ryder
1967).
Whitefaced Ibises feed actively by Walking, slowly to
quickly, while Probing rapidly, usually at each step. Period-
ically they stop and Stand while Probing at the base of
aquatic plants. They also Head Swing while standing still in
open water, vibrating their mandibles rapidly (Belknap
1957).
The species is highly gregarious, probably more so than
the Glossy Ibis. Flocks can exceed 1000 birds (Belton 1984).
Flocking, feeding birds are often reported to engage in Leap-
frog-feeding, in which the rear birds of the flock hop to the
head of the moving flock. Birds in the flock are often sub-
jected to piracy by other birds, such as grackles (Quiscalus
spp.). Birds fly to and from feeding areas and roosts in
flocks, and they roost together in trees or on the ground in
marshes.
They feed on small fish, frogs (Hyla), crayfish, snails,
bivalves, earthworms and aquatic insects such as whirligig
beetles, beetle larvae, midge fly larvae and pupae and
dragonfly nymphs (Belknap 1957, Centro Editor de Amer-
ica Latina 1983a). In Nevada and Utah, they feed primarily
on earthworms but also on dragonfly larvae, horsefly larvae,
beetles caterpillars, and other insects, snails and spiders
(R.A. Ryder 1967, Bray and Klebenow 1988). In Argentina
they feed with gulls on flies and fly larvae associated with
livestock carrion (Weller 1967).
BREEDING
Whitefaced Ibises are highly conspecifically gregarious,
nesting in large numbers in compact colonies, usually in
marshes. This species preferentially nests in herbaceous
marsh vegetation, although it will nest on short bushes, on
dry land and on spoil islands. The most common nesting
habitat throughout its range is tule marsh (Scirpus spp.), as
well as common cane (Phragmites) inland and cordgrass
(Spartina spp.) along the coast. In North America, it is likely
that the use of high ground on spoil islands may increase as
natural marshes become less suitable.
This species changes its nesting locations frequently,
causing considerable year-to-year shifts in breeding areas.
In part, such shifts are due to changing availability of
nesting and feeding habitat in flooded shallow marshes and
wet meadows. It is likely that birds return to previously
used nesting sites, but if conditions are unfavourable begin
to search for more appropriate conditions, becoming
attracted to waterbird colony sites elsewhere. This sugges-
172 Whitefaced Ibis
tion is supported by the fact that colonies are unsynchron-
ized, suggesting independent arrival of breeders.
Typically, the Whitefaced Ibis nests in the spring, April
to May in North America, and November to December in
South America (Belton 1984, S. Narosky pers. comm. 1990).
However, they can delay breeding, for example to mid June
in the USA, if conditions are adverse (Belknap 1957), or
may even skip years in nesting. Because nesting is often not
highly synchronized, large chicks and eggs may be present
at the same time.
Early courtship has not been recorded in detail. The male
displays from several potential nest-sites, especially from
abandoned nest-platforms. He may also attempt to displace
other birds from their platforms using Bill Thrusts and
Supplanting Flights (Belknap 1957). At the display site, the
male uses a ritualized bill probing, and gives a 'Hunk-hunk-
hunk' call, which the female answers. At a potential nest-
site both birds preen, perhaps Display Preening. The actual
nest-site selection is by the female (Belknap 1957).
The nest is built on top of grasses and sedges, which are
bent over to support the structure. The birds tend to nest
relatively low, within 1 m of the water, even when in shrubs.
Often, however, the nests end up floating on the water
surface or on the ground. In Texas, 86% of nests were found
to be on the ground, the rest in the branches of low shrubs
(Burger and Miller 1977).
After pairing, the male and female spend considerable
time preening, perhaps Display Preening, and Allopreening.
Initially the female gathers reeds for the nest, which the
male guards. However, after construction or renovation
begins in earnest, the male gathers while the female con-
structs the nest. Copulation occurs at the nest. Nest
exchange is accompanied by mutual 'Geeeek, geeeek5 calls
(Belknap 1957).
Usually, although not always, the nest is relatively large:
40 cm across, solid and flat, but with a well-developed cup.
Nests are usually made of reeds or other herbaceous vege-
tation, sometimes with sticks included. The cup is lined with
shredded bits of plant material such as reed leaves. White-
faced Ibises also frequently take over previously built nests
of their own or other species, adding to and reconstructing
them to meet their own needs.
Burger and Miller (1977) found them to be more tolerant
of other Whitefaced Ibises near their nest than they were of
other species. None the less, they are extremely defensive of
their nest and an area of up to 1m around it. The bird
defends with Bill Thrusts and Supplanting Attacks,
occasionally landing on an opponent's back, pulling its
feathers or striking blows with its bill, while giving 'Gheeee'
calls.
Eggs are laid at 1-2 day intervals and hatch asynchro-
nously after 20-22 days' incubation. The clutch size is 2-5,
averaging 2.6 in Texas, 2.7 in Utah, 3.0 in Louisiana (Bel-
knap 1957, Kotter 1970, K.A. King et al. 1980). The eggs
are deep blue to greenish blue lacking any mottling. Utah
eggs averaged 52.0 X 36.7 mm and 37.6 g.
The young call with a vibrating 'Cheeeeeeeeu' and are fed
by regurgitation (Belknap 1957). Parents arriving to feed
young give a 'Goick, goick' call. Eggs and young often suffer
heavy losses, especially from heat and predation (Belknap
1957). In Utah, 66% of eggs hatched and 53% fledged.
After fledging, young fly with adults to foraging areas,
returning at dusk to roost. The young birds can fly in their
fifth week and are independent in 8 weeks. First-year mor-
tality is estimated to be 54%.
TAXONOMY
The Whitefaced Ibis is closely related to the Glossy and
Puna Ibises. Palmer (1962) considered the two North Amer-
ican forms to be conspecific. Nesting areas overlap slightly
in southern United States, and the American Ornitholo-
gist's Union (1983) considers the two forms to be separate
species, but part of a superspecies. The specific name has
previously been given as guarauna and as mexicanus, but these
properly refer to other species (Parkes 1955).
CONSERVATION
The general decline and contraction of breeding colonies in
the 1960s was caused by pesticide contamination. High
pesticide levels in various populations have been shown to
affect clutch size, eggshell thickness and other aspects of
reproductive physiology (Flickinger and Meeker 1972,
Capen 1977a, 1978, K.A. King et al. 1978, Capen and
Leiker 1979, K.A. King et al. 1980, Steele 1984). Population
declines have occured in Nevada, Utah and Texas (R.A.
Ryder 1967). In Texas, the population decreased by 77%
from 1969 to 1974 because of pesticides (K.A. King et al.
1980). Despite discontinuation of the use of aldrin in rice
fields in the USA, pesticides are continuing to be accumu-
lated by Whitefaced Ibises wintering in Mexico (R.A. Ryder
1985, Hennyetal. 1985).
In many areas, ibises are used for food, and this species is
no exception. An open hunting season existed for White-
faced Ibis in California until 1915 (Bent 1926). They remain
hunted in Mexico and in areas of South America. Chilean
populations have apparently been persecuted in the past,
wherever colonies were accessible to poachers (Schlatter
pers. comm. 1990). Such hunting would be particularly
troublesome in areas that are otherwise marginal, such as
Chile.
In recent years, the species has expanded its range in
North America, with nesting records occurring over all of its
former range in the USA in places where it had been absent
for decades, (e.g. Minnesota, Idaho, New Mexico, and
California) and even over an expanded area. In Oregon,
numbers have increased from 600 to 2595 pairs during
1980-1987 alone (Ivey et al. 1988). This expansion co-
incides with a change in pesticide use as well as creation of
aquatic habitats by dams and irrigation.
Additional information on the status of various popu-
lations is needed. The most critical concerns are a re-
duction, throughout the Whitefaced Ibis' range, in the use
of persistent pesticides that adversely effect them and the
protection of aquatic habitats, especially marshes.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 309.
 Puna Ibis
Plegadis ridgwayi (Allen)

Falcinellus Ridgwayi J.A. Allen, 1876, Bull. Mus. Comp. ZooL, 3, p. 355: Lake Titicaca, Peru

Other names:

Andean Glossy Ibis (English); Cuervo de Pantano de la Puna, Cuervo de Pantano del Norte,
Cuervillo de la Puna, Cuervillo de Bolivia (Spanish); Make-make, Yanahuiku (South American
Indian)

IDENTIFICATION
The Puna Ibis is a medium-sized (56-61 cm tall) dark ibis, looking black from a distance, with a dark
red downcurved bill.
The dark purplish brown base colour of the adult's body is highlighted by deep glossy purple and
green reflections above on the lesser wing-coverts, becoming almost black below. The head and neck are
dull chestnut. The wings and tail are glossy green, the tail a darker green. The iris is red; the legs and
feet are black. In winter, overall coloration is duller, and the head and neck become darker brown with
fine white streaking.
During the breeding season, the head and neck turn deep chestnut. The species is sexually dimorphic,
in that males are considerably larger than females. In 10 males, the culmen averaged 11.7 cm (10.8-
12.7 cm), whereas in 7 females it averaged 9.2 cm (8.6-9.5 cm) (Blake 1977).
Chicks are dark brown with banded bills. The juvenile is similar to the non-breeding adult, except
that it has more conspicuous white streaks on the neck, extending onto the belly, and lacks much of the
adult's sheen. Its iris is brown and its bill lighter and more brownish.
The Puna Ibis flies in flocks, sometimes large ones, forming Vs or wavy lines, usually low over the
terrain. It is usually silent, but gives 'Wut' and 'Cwurk' calls when disturbed (Fjeldsa and Krabbe
1990). Juveniles give a whispering call.
The Puna Ibis is a glossy ibis, although larger than the Whitefaced Ibis. It is also duller and stockier
than the Whitefaced, with shorter legs and lacks the band of white feathers around the bill. It is darker
and longer-legged than the co-occurring Buffnecked, and does not have the stocky, broad-winged and
short-tailed appearance of that species (J. Fjeldsa pers. comm. 1990).

DISTRIBUTION AND POPULATION

The Puna Ibis is endemic to the Andean Mountains of western South America above 3500 m. It occurs
in north-central Peru (Ancash and Junin) southward to the Titicaca basin of Peru and Bolivia, to
northern Chile (Arica, Tarapaca). It also occurs very locally in Cochabamba, Bolivia and in north-
western Argentina (Jujuy, Salta) (Blake 1977, Fjeldsa and Krabbe 1990).
It is a wanderer and is seen regularly as high as 4800 m above sea level in Peru (J. Fjeldsa pers.
comm. 1990). It winters, from May to September, beyond its breeding range as far as valleys at the foot
of the Andes, along the Peruvian coast (D.L. Pearson and M.A. Plenge 1974, Hughes 1984, Fjeldsa and
Krabbe 1990). It is accidental in La Libertad, north Peru, but it is likely that reports from Central
America and Patagonia are misidentifications (J. Fjeldsa pers. comm. 1990).
Little is known about population sizes but, overall, they may be considered common. The species can
be seen at most Peruvian puna lakes, though usually only in small numbers. However, it can be locally

abundant, and in the Lake Junin area of Peru, the popu-
lation has been estimated to be about 8000 birds (Douro-
jeanni et al. 1968). However, it may nest only in a dozen or
so of the puna lakes of the Andes (J. Fjeldsa pers. comm.
1990).
ECOLOGY
This species is characteristic of the high-altitude puna, the
temperate zone of the Andes. It may even be found in the
higher arctic zones, but it seldom occurs in the subtropical
zones except on the Peruvian shores (J. Fjeldsa pers. comm.
1990).
Puna Ibises frequent large marshes and damp pasture-
land, where they forage on muddy flats, along creeks and in
short grass (Fjeldsa and Krabbe 1990). They also feed
terrestrially, in grasslands quite far from water. They
appear to be rather opportunistic, congregating, for
example, in places having large numbers of brightly
coloured lepidoptera larvae (J. Fjeldsa pers. comm. 1990).
The typical feeding behaviour is Probing deeply in the
water or mud to extract prey, usually along the edges of
denser vegetation (J. Fjeldsa pers. comm. 1990). As is usual
Puna Ibis 175
for Plegadis ibises, this species feeds socially, sometimes in
flocks containing large numbers of birds.
BREEDING
The Puna Ibis nests in tall reeds in highland marshes. In the
enormous reed-beds of Lake Junin, it locates its nesting
colonies in places that are quite inaccessible to humans (J.
Fjeldsa 1983, pers. comm. 1990). The birds especially use
very tall Scirpus reed beds, isolated by trenches of deep
water. In other areas, such as on the Puno—Cuzco border,
they nest in Juncus-lintd ponds. They may nest in mixed
colonies with other waterbirds.
The nesting season is somewhat variable geographically
(Fjeldsa 1983, pers. comm. 1990, Fjeldsa and Krabbe 1990).
At Lake Junin, Peru, nesting is in April to July, with some
clutches also in January and February. Nesting is in Janu-
ary to March in Ayacucho, and in November to January
and February in the southern parts of the range.
Nothing is known about the species' courtship behaviour.
CONSERVATION
The biology of the Puna Ibis is almost entirely unknown,
and its status throughout the Andes is not well understood.
It historically inhabited a large portion of the puna zone,
but it is now considered vulnerable in Chile (Glade 1988).
Fortunately, a reasonable recovery of the local population
has occurred in Lauca National Park (Schlatter pers. comm.
1990).
Much of the marshy vegetation of the puna zone, which
supports the ibis, is under threat (Fjeldsa 1985a). A combi-
nation of human overpopulation, harvesting of tules and
cattle grazing have opened much of the marsh. J. Fjeldsa
(pers. comm. 1990) reports that some clearing of tules, as
has occurred along Lake Titicaca, may be advantageous
because it creates feeding habitat for the ibis. However, it
becomes harmful when safe nesting areas are eliminated. In
many parts of the high Andes, grazing by cattle has com-
pletely eliminated all the tall marsh vegetation. It has been
suggested that the rarity of ibises in the Bolivian highlands
is due to egg-collecting traditions of the Ayumara Indians
(Fjeldsa and Krabbe 1990).
Lake Junin, one of the primary nesting sites for the Puna
Ibis, remains one of the most productive waterbird habitats
in the Andes. Proper conservation and management of the
lakes and wetlands of the puna zone are essential to the
preservation of this endemic ibis.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 309.
 Sharptailed Ibis
Cercibis oxycerca (Spix)

Ibis oxycercus Spix, 1825, Avium Species Novae Itinere Brasiliam, 2, p. 69, pi. 87: in Provincia Para

Other names:

Longtailed Ibis (English); Tarotaro (Spanish); Trombeteiro, Tara (Portuguese); Groote Glans-
Ibissen (Dutch)

IDENTIFICATION
The Sharptailed Ibis is a dark, medium-sized ibis, c. 75-86 cm long, having a conspicuously long tail
that has suggested its common English name.
The adult's plumage is black overall, usually showing a green glossing. The upper back, wings and
tail have a purplish gloss. A fuzzy occipital crest is often noticeable, running along the back of its head
and upper neck.
The bill, legs, toes, and bare face are orangish red. The throat is yellow-orange, and a feathered strip
of grey extends from the lower bill below the eye. The iris is greyish red, but Betsy Thomas (pers.
comm. 1990) has observed a bird with scarlet coloured irides; this colour may be associated with
breeding. Additional information on soft-part colours, especially during courtship, is needed.
The sexes appear to be similar. Although the sizes of museum specimens labelled as to sex overlap, in
the wild we have observed what appears to be a size difference between members of a pair, suggestive of
sexual differences in size (pers. obs. J.A.K.).
Hatchlings appear not to have been carefully described. Juveniles are similar to adults but lack a
metallic sheen to their feathers.
This is a noisy bird, which has attracted our attention from a distance by its calls, given while
perched in trees. Its usual vocalization is a single or doubled 'Cuk' or 'Tur-cuck' that can blend with a
bugled 'Tur-dee' call, which gives this ibis its Spanish common name, Tarotaro. The call is similar to
but less nasal and raucous than that of the Buffnecked Ibis (Sick 1985, W. Belton pers. comm. 1990).
The flight call is a 'Cuk-cuk-cuk-cuk'.
In flight the Sharptailed Ibis is relatively slow, and flying appears somewhat laboured. When a bird
flies up, it usually goes but a short distance, usually to a nearby tree.
This is a quite distinctive ibis, rather larger and stockier than sympatric species. When at rest, its
long tail projects well beyond the tips of its folded wings, and, when in flight, beyond the trailing legs
and feet. It is larger than the other dark ibises, such as the superficially similar Glossy Ibis, and can be
further identified by its reddish face and soft parts.

DISTRIBUTION AND POPULATION

The Sharptailed Ibis is a lowland bird of the Amazon and Orinoco River basins. It occurs from east of
the Andes in Colombia through inland Venezuela, following the Orinoco and Apure rivers to the coast,
and again in a disjunct distribution, along the inland portion of the northern Amazon drainage from
southwestern Guyana, to northwestern Brazil (Rio Negro), western Brazil (Rio Guapore), and south to
northwestern Mato Grosso.
The species is very locally distributed within its overall range. Although probably not substantially
migratory, it does undertake seasonal population shifts. Both E. Aguilera (pers. comm. 1990) and
178 Sharptailed Ibis
B.T. Thomas (1979a) found it to be absent from the interior
llanos during the dry season (December, January). It can be
fairly common and obvious locally, but nothing is known
about population sizes. More detailed information on range
and population is much to be desired.
ECOLOGY
This is an ibis of llanos savannas and river-edges of Amazo-
nia, below 300-500 m. During studies in the Venezuelan
llanos, we found it to be more terrestrial than the flocking
ibises, feeding in relatively short grass and open areas in
relatively higher and drier (but still moist) ground (pers.
obs. J.A.K.). It generally feeds in water less than 3 cm deep,
but it also frequents pools and muddy spots along the edges
of ponds and streams in open areas, in short grass, and in
shallow water (Kushlan et al. 1985).
Food finding seems to be primarily tactile. It usually
forages by Walking rather quickly while slowly Probing
deeply into the soft substrate. B.T. Thomas (pers. comm.
1990) has observed a pair feeding on dry ground by tossing
leaves aside before probing in the soft sand underneath.
Sharptailed Ibises do not flock, and seldom are seen
alone. Rather they typically feed in pairs or in very small
groups, usually including 2 adult birds (parents) and
fledged young. These young may Beg food from the adults.
Little is known about the diet. They appear to eat inverte-
brates. More information is needed on this point.
BREEDING
The Sharptailed Ibis nests solitarily. The nests are placed in
trees in gallery forests along streams and in isolated cop-
pices within the grassland.
Birds appear to remain paired year-round, and courtship
is probably rather perfunctory. In any case, we know of no
information on courtship behaviour. Mutual preening
occurs between members of a pair (Ogden and Thomas
1985b).
In the llanos, E. Aguilera (pers. comm. 1990) found
nesting to begin in August and September and to continue
through February. This is in the drying season, suggesting a
dependence on exposed higher ground for effective foraging.
TAXONOMY
The Sharptailed Ibis is generally similar to the other dark
ibises, but its taxonomic relationships remain to be dis-
covered through careful study.
CONSERVATION
Basic to even considering the conservation of this species is
our limited knowledge. Little is known about its basic bi-
ology and ecology, and such information is sorely needed.
Because it is a solitary bird, its status may decline without
anyone's knowledge (C.S. Luthin pers. comm. 1990).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 310.
 Plumbeous Ibis
Harpiprion caerulescens (Vieillot)

Ibis caerulescens Vieillot, 1817, Nouv. Diet. Hist. Nat., nouv. ed., 16, p. 18; based on 'Curucau
aplomado', no 363, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p. 195:
Paraguay

Other names:

Blue Ibis (English); Curicaca, Curucau Moroti, Bandurria, Bandurria Mora, Vanduria (Spanish);
Macarico-real, Curicaca, Guricaca-cinza (Portuguese)

IDENTIFICATION
The Plumbeous Ibis is a relatively large grey ibis (71-77 cm long), identifiable especially because of a
shaggy crest along the back of its head and neck and a distinctive white bar on its forehead.
The basic plumage colour of the adult is grey to bluish-grey, lighter below than above because the
dorsal feathers are each broadly mottled with greyish brown. The head is almost entirely feathered,
with a very noticeable white band of feathers on the forehead and a less noticeable smaller band at the
lower side of the bill. The distinctive nuchal crest, extending a full 10 cm down the back of the neck, is
composed of long, grey plumes each having a cream-coloured centre, which in some lights gives the
neck a pinkish tinge. These are arranged so that they appear furrowed (Sick 1985, W. Belton pers.
comm. 1990). The flight and tail feathers are dark brown to black, glossed with a greenish bronze that
can look silvery in the appropriate light.
The strong, downcurved, distinctly serrated bill is black. The exposed skin of the lores and chin are
grey to black. The legs are a pinkish orange, which brightens in the courtship period (F. Olmos pers.
comm. 1990). The iris colour is orangish and has been variously reported as brownish yellow, orange-
yellow and orange (Naumburg 1930, Blake 1977, Belton 1984). Wetmore (1926:63) noted that the iris
was somewhat lighter on its inner margin. He also noticed in one specimen that its lower eyelid was
coloured lilac.
Based on our examination of museum specimens the species appears to be sexually dimorphic, with
males larger than feinales on average (pers. obs. J.A.K.).
Nestlings have a light grey down, reddish skin, a white forehead and a black bill (Cintra 1986).
Young have a less noticeable crest, and the white forehead band seems broader (F. Olmos pers. comm.
1990).
The Plumbeous Ibis flies with strong, steady flapping with head and neck extended, frequently
gliding.
The calls are well known. Hudson (1920:123) likened its voice to a dozen strokes on the brass strings
of a banjo so big that it could be heard a mile and a half away. He called it the voice of the desolate
marshes. Belton (1984:433, Sick in prep, as transliterated by W. Belton pers. comm. 1990) have
described its vocalizations. The basic call is a rapid, high, loud Ti-pi-pi-pi'. The flight call is a powerful,
distinctive, trumpeting, which has been rendered as 'Kree-kree-kree', 'K-k-kuh-kuh-kee-kee-kee-kee-
keh-kuh-kuh', and 'Ki-ki-ki, Ghew-ghew-ghew'. These distinctive calls are an indication of the need for
long-range communication, particularly between mates (Remsen 1986).
The Plumbeous Ibis is rather distinctive among sympatric ibises owing to its thick neck and solid
coloration. The Buffhecked Ibis is found in many of the same areas but differs in coloration (F. Olmos
pers. comm. 1990, W. Belton pers. comm. 1990).

DISTRIBUTION AND POPULATION
The Plumbeous Ibis is a bird of the great marshy grasslands
or pantanal of south-central South America. It occurs from
northern and eastern Bolivia east to Beni, Brazil, in central
Mato Grosso and in Rio Grande do Sul, Paraguay, Uruguay
and Argentina, including Tucuman, Formosa, Santiago del
Estero and northern Santa Fe. It is casual south in Argen-
tina to Cordoba and Buenos Aires Province.
All indications are that individuals appear to remain in a
specific area year round (F. Olmos pers. comm. 1990).
However, they are occasionally found outside the normal
range and earlier reports spoke of some seasonality to move-
ments, so additional detailed information is desirable on any
short-range seasonal or dispersive shifts that might occur.
This species is locally well known, but numerical infor-
mation on population sizes are lacking. It is fairly common
in some areas, such as the northern pantanal of Pecone,
Mato Grosso, Brazil (F. Olmos pers. comm. 1990). But it
has, overall, a rather patchy distribution, occurring regu-
larly in some locations but more rarely in others (Wetmore
1926:63, Belton 1984:433).
ECOLOGY
The Plumbeous Ibis is a bird of wetlands, especially the
seasonal marshes of the chaco and pantanal. It feeds in
ponds, marshes, swamps, flooded pasture and other moist
ground, especially drying pools of water. Although usually
occurring in wet areas, it may also be encountered on drier
grasslands and is commonly found in association with hu-
mans. It roosts in trees near water and is often seen resting
in such locations.
Plumbeous Ibis 181
Usually the foraging method is Walking slowly, Probing
in the mud and shallow water. Birds frequently Probe rather
deeply, completely submerging their bills in the process.
This is a solitary species, found singly or in pairs and
more rarely in small groups of 4-6 birds. It would seem that
the larger groups include members of a mated pair, season-
ally accompanied by fledged juveniles (F. Olmos pers.
comm. 1990). It also may feed in company with Whitefaced
Ibises (Sick 1985:179). Brooks (1991) reported a flock of
more than 40 birds, but it does not form large conspecific
flocks as do some ibis species.
It is known to feed on snails (Pomacea), crabs, aquatic
insects, frogs, fish, and eels (Sick 1985:179, Cintra 1986,
R.W. Storer 1989). Further details of its diet are needed.
BREEDING
The Plumbeous Ibis nests solitarily within sparsely wooded
areas (Dubs 1988), As it is very likely that these birds
remain paired year round, the nest is probably placed
within the usual home range of the parents.
In Mato Grosso, nesting begins in March and April and
continues at least until mid-October (Dubs 1988, Olmos
1990, Cintra 1986). This timing corresponds with the drying
season, with fledging occurring towards the end of the dry
period.
Both members of the pair construct the nest, which is
made of twigs and lined with leaves and grass (Dubs 1988).
Nests are about 50 cm wide and 25 cm high (Cintra 1986).
They are placed on horizontal limbs of the large trees; nests
in the pantanal of Mato Grosso are reported to be up to
12 m above ground (Cintra 1986, Dubs 1988).
Clutch size is 2-3 in the pantanal. The eggs are pale grey.
Incubation is 28 days, with young hatching in early Sep-
tember (Cintra 1986, Dubs 1988).
The young are heavily guarded by one of the parents
during the first 10 days. When approached by humans, the
ibis screams loudly, lowers its body, bristles its feathers and
jabs at the intruder with its bill. The young also defend
themselves with a forward lunge of the bill (Cintra 1986).
These behaviours suggest that the risk of predation is high.
Olmos (1990) described predation on nest contents by mon-
keys and a hawk.
Nests in the pantanal have 2 young that fledge at 40-
43 days (Cintra 1986, Dubs 1988, F. Olmos pers. comm.
1990). They initially feed in marshes near the nest and then
move away to feed with the family, returning to the nest-tree
to sleep for several weeks (Cintra 1986).
TAXONOMY
The Plumbeous Ibis has long been considered to be a very
distinctive ibis, having been variously placed in several
genera over the years, including Harpiprion, Molydophanes,
Geronticus and, more recently, Theristicus, the latter sugges-
ting a relationship to the BufFnecked Ibises. Short
(1975:196) retained the genus Harpiprion, considering this
species to have no close relatives. Until additional research
182 Plumbeous Ibis
is conducted into its taxonomic relationships, it seems best
to retain the species in its traditional monotypic genus.
CONSERVATION
As is the case for some of the other South American ibises,
so little is known about this species that neither its biology
nor its status is sufficiently understood. Such information is
essential if its conservation is to be assured.
Fabio Olmos (pers. comm. 1990) reports that the species
is not persecuted in the pantanal of Mato Grosso and is
often associated with humans. As such, it is vulnerable to
the burning and logging of nesting trees, which are found on
the higher ground also used for farming. Human association
may also place them at risk from pesticides, especially in
heavily cultivated regions such as the Taquari Basin.
Although the pantanal of Mato Grosso remains relatively
remote, future threats, such as hydroelectric dams, mining
and expansion of agriculture, will also adversely affect habi-
tat protection (D.A. Scott and M. Carbonell 1986:91).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 310.
 Buffnecked Ibis
Theristicus caudatus (Boddaert)

Subspecies:
Theristicus caudatus caudatus (Boddaert)
Scolopax caudatus Boddaert, 1783, Table Planches Enlum, p. 57; based on 'Courley a col blanc', de
Cayenne of Daubenton, 1765-81, Planches Enlum, pi. 976: Cayenne, French Guiana
Theristicus caudatus hyperorius Todd
Theristicus caudatus hyperorius Todd, 1948, Proc. Biol. Soc. Washington, 61, p. 50: Buena Vista,
Bolivia
Theristicus caudatus melanopis (Gmelin)
Tantalus melanopis Gmelin, 1789, Syst. Nat., 1, p. 653; based on 'Black-faced Ibis5 of Latham, 1785m
General Synop. Birds, 3, p. 108, pi. 79: 'In insula novi annf = New Year's Island, near Staten
Island, ex Latham
Theristicus caudatus branickii Berlepsch and Stolzmann
Theristicus branickii Berlepsch and Stolzmann, 1894, Ibis, p. 404: Maraynioc, Pariayacu, Junin,
Peru

Other names:
Whitenecked Ibis, Blackfaced Ibis, Whitethroated Ibis, Andean Ibis, Grey Ibis (English); Bandurria,
Bandurria Amarilla, Bandurria Comun, Bandurria del Norte, Bandurria del Sud, Bandurria de
Verano, Bandurria de la Palma, Vandurria de Invierno, Mandurria, Tautaco, Curacau, Tutachi,
Carcel, Code, Corococco, Gangola, Garza Pelata (Spanish); Curicaca, Despertador (Portuguese); Le
Petit Courly d'Amerique (French); Witkin-Glans-Ibissen (Dutch)

IDENTIFICATION
The Buffnecked Ibis is a relatively large light-coloured ibis, 71-81 cm long, standing c. 43 cm tall. It is
distinguished by its distinctively buffy white neck and head and, in flight, by a large white patch
contrasting with dark wing feathers.
The basic colour pattern of the adult is rather complex and also variable among tropical, temperate
and montane populations. The uppersides are basically grey highlighted by a slight greenish gloss and
light grey to white wing-coverts, which show as the wing-patch noted above. The cream to buff colour of
the head and neck, which is set off from the body coloration, turns a deeper yellow during courtship,
fading to a whiter shade afterwards, probably due to wear. The top of the head is cinnamon rufous, as
often are feathers at the base of the neck. The undersides are paler or darker than the base colour,
depending on the population; the rump and undertail are black. A distinctive pectoral band of grey
feathers crosses the chest in light-bellied forms. The grey flight feathers have a greenish gloss.
The two northern subspecies, caudatus and hyperorius, have dark undersides, whereas the Andean and
southern subspecies, branickii and melanopis, have light undersides with pale colour on the breast
reaching further down on the belly although interrupted by the grey pectoral bar (Fjeldsa and Krabbe
1990, J. Fjeldsa pers. comm. 1990). It is not certain that the trends of variation in the species are
appropriately captured within the definitions of the four subspecies, as there is much intergradation
among the typical forms, and examples exist of birds outside the defined range of their plumage type
(Friedmann and Smith 1950). Much of the variation is probably clinal, that is, associated with climatic
differences from tropical environments to cooler and high-altitude environments, which tend to be

occupied by smaller and lighter plumage birds. Essentially,
the undersides become paler going from north to south and
from low to high altitudes.
The northern tropical subspecies, caudatus, has the dark-
est, blackish, underparts. Its neck is whitish buff with a
rufous patch below. The wing-feathers have some white on
them, especially on the coverts and parts of the primary
feathers.
The southern tropical subspecies, hyperorius^ is very simi-
lar to caudatus, and may not be well separable. It has lighter
but still relatively dark undersides, grading from brownish
to grey-brown. Its dorsal colour is also a bit browner than
caudatus. The grey feathers have a lighter grey edging, mak-
ing them more conspicuous (Todd 1948). Its tail has a green
sheen. These birds generally have larger bills than caudatus
(C.S. Luthin pers. comm. 1990).
The southern temperate melanopis, has a richly buff fore-
neck and breast and a rather restricted throat feathering,
sometimes limited to the chin alone, giving it the fullest
gular pouch of the recognized subspecies. The wing-coverts
are pale grey to amost silvery white.
The Andean subspecies, branickii, is somewhat smaller
than the other subspecies but has the longest wings (C.S.
Luthin pers. comm. 1990). The base of the neck is tinged
with rufous. It has the palest breast and undersides, which
are pale buff to creamy white. Its wing-coverts are a rather
dark grey barred with paler feather-tips showing as a grey-
silver (J. Fjeldsa pers. comm. 1990, C.S. Luthin pers.
comm. 1990). Overall, its dorsal coloration is more consist-
ently grey than in the other forms.
In all subspecies, the skin around the eye and on the sides
of the throat or entire throat is black, and may be somewhat
Buffhecked Ibis 185
warty. The dark face has inspired one of the species' com-
mon English names, the Blackfaced Ibis. The iris is brown-
ish red, often described as being red to crimson. The legs
and feet are pink, orange-red or purple-red. The chin retains
a thin line of feathering, less in melanopis.
Along with the plumage, the soft parts brighten during
the breeding season. The legs turn redder at the start of
courtship. They have also been reported as black tinged
with crimson (P.S. Humphrey et al. 1970:104). Both males
and females possess an expanded gular sac during this time.
The sexes are similar in plumage, but males are larger in
body measurements and weight. In Chilean populations,
males averaged 1758 g, and females averaged 1598 g (Gantz
in R. Schlatter pers. comm. 1990).
At hatching, the young have little feathering, but a grey
down soon develops. J. Fjeldsa (pers. comm. 1990) reported
that a young bird he collected in Patagonia was uniformly
grey, but Blaauw (1917) reported that the head and breast
of nestlings are yellow-white. The legs are initially light
pink, the throat blue-black and the bill blue-white with
black at the tip. The legs then turn grey, with a purple tinge.
The iris is brown; the facial skin is black, with a white patch
on either side at the base of the bill.
Juveniles are similar to adults, but their buff colour is
duller and their abdomens are paler than in the adult. The
head and neck are streaked brownish. The wing-coverts
have buff edges, giving a scalloped effect (Fjeldsa and
Krabbe 1990). The head turns more rufous with age. Soft
parts are duller than in the adult. The legs are pink and the
iris is dark brown (B.T. Thomas pers. comm. 1990, Schlat-
ter pers. comm. 1990). An immature bird in the American
Museum of Natural History had a greenish bill and brown-
flesh feet (P.S. Humphrey et al. 1970:104).
In feeding and flying, the Buffnecked Ibis is quite vocifer-
ous, frequently giving its distinctive two-note call. On the
ground, this call may be given with the head thrown back;
in flight, only the first syllable may be called. It is a musical,
bell- or trumpet-like call that has been variously rendered
because it sounds different depending on distance and ter-
rain. In English, it may be rendered as 'Tur-tac*, accenting
the second syllable. It is better described as being reminis-
cent of a blacksmith's hammer on an anvil. Hudson (1920)
described the call of melanopis as short, abrupt notes of
extraordinary power resembling the blows of a powerful
hammer on a metal plate. In 1860, Darwin described the
call as being like the neighing of the wild llama (the gua-
naco) (Darwin 1962:166).
The Buffnecked ibis flies with great dexterity on quick,
strong wing-beats. In flight, the legs do not extend beyond
the tail, and the light wing patches flash conspicuously. The
birds often fly low over to the ground but sometimes soar to
great height during their daily wanderings (J. Fjeldsa pers.
comm. 1990), and they characteristically undertake com-
plex aerial manoeuvres. Their acrobatics going into roost
have been much remarked upon. Hudson (1920) described
the performance as frenzied, the birds simultaneously dash-
ing downward with 'a violence wonderful to see'. Indi-
viduals and flocks, where they occur, often wheel and zigzag
while calling loudly. The birds rise and fall, sometimes
186 Buffnecked Ibis
together and sometimes scattered, while calling with their
'loud percussive screams' (Hudson 1920).
The species should not be easily confused with any other
ibis, owing to its distinctive call, neck colouring and, in
some races, the bicoloured undersides. The Plumbeous Ibis
is monocoloured and has a thicker neck. The Buffnecked
Ibis might look like a goose from a distance, but not up
close.
DISTRIBUTION AND POPULATION
The Buffnecked Ibis occurs over much of South America,
from Colombia southward. Hiatuses in distribution occur,
such as the highlands of southern Colombia, southern Vene-
zuela and Brazil, parts of the Ecuadorian and Peruvian
Andes, the coastal desert of Chile, inland northern Chile,
and western Argentina (Zapata and Martinez 1972, Blake
1977, Fjeldsa 1985a, Fjeldsa and Krabbe 1990, J. Fjeldsa
pers. comm. 1990, Luthin pers. comm. 1990). It is acciden-
tal in Panama to the north and the Falklands to the south-
east.
Theristicus caudatus caudatus is the northern lowland form,
ranging from Colombia south to Valle del Cauca and Meta
and Venezuela south to Apure and Bolivar, to the Guianas,
and into northern Brazil. It is this race that is casual in
Panama (Pacora and San Jose) (Blake 1977:194).
Theristicus caudatus hyperorius occurs in central and southern
Brazil, eastern Bolivia, northern Argentina (Tucuman,
Chaco, Santa Fe, Entre Rios), Paraguay, southeastern Bra-
zil and Uruguay, with an isolated small population above
2000 m in the Cordoba Mountains near Cordoba, Argentina
(Arballo 1990, J. Fjeldsa pers. comm. 1990).
Theristicus caudatus branickii is the northern highland sub-
species, occurring very locally in the mountains of Ecuador
(Antisana, Cotopaxi), Peru (Ancash, Junin, Huancavelica,
Ayacucho, Cuzco, Puno), northwestern Bolivia (La Paz,
Oruro), and extreme northern Chile (Arica), rarely moving
down to the lowland valleys of Chile, Peru and Bolivia
(Cochabamba) (Fjeldsa and Krabbe 1990).
Theristicus caudatus melanopis is the Patagonian subspecies
(Blake 1977, Fjeldsa and Krabbe 1990, J. Fjeldsa pers.
comm. 1990). To the north, it occurs in relict populations
along the Andes in Peru (Lambayeque, La Libertad, Lima,
Tacna). It is absent from the Atacama Desert, occurring
again south of Antofagasta, then down the coast to Tierra
del Fuego, including the offshore islands. In the northern-
most part of this range, it occurs in the high mountains,
whereas further south it uses the central valley of the Andes
and the coastal zone, and even further south it occurs across
the whole of Chile (A.W. Johnson 1965). In the southern
Argentine, the range extends northward through the Pata-
gonian grasslands to Neuquen and Rio Negro.
Many populations appear to occur in the same area year-
round. However, the southern populations are migratory,
and other birds may shift seasonally, especially in the cool
or dry seasons. The southern melanopis populations move
northward to Chile and Argentina as far as Tucuman,
Cordoba and Buenos Aires Province, the latter destination
being the main wintering ground for Patagonian birds
(MacDonagh 1942, Nores and Yzurieta 1980, J. Fjeldsa
pers. comm. 1990). They may be more common there in dry
years (Weller 1967). This subspecies is also the one that is
accidental in the Falklands. The movement of southern
populations to the north in the winter is the origin of several
of southern South American common names, such as Ban-
durria de Verano.
Although the northern populations of the llanos remain
year-round, some movements apparently take place, as they
are less abundant in the dry season (E. Aguilera pers.
comm. 1990). Similarly, Andean birds may shift to lower or
wetter locations, including into the pampas of Argentina,
during the winter dry months (Fjeldsa and Krabbe 1990,
C.S. Luthin pers. comm. 1990).
Little is known about population sizes, undoubtedly be-
cause of the birds' wide dispersal. Overall, however, where
they occur, most populations appear to be relatively high.
The species seems particularly common in the grasslands of
the llanos and Patagonia. Montane populations of branickii
have always been considered to be rare, perhaps because of
their highly localized distribution, their remoteness and
their wary habits (Fjeldsa 1988, Fjeldsa and Krabbe 1990,
C.S. Luthin pers. comm. 1990). It appears that this species
is fairly common in the southern Andes but much rarer
further north. More information is needed on the status of
this species in the Andes.
ECOLOGY
The Buffnecked Ibis is a bird of the damp to dry grasslands
and meadows of South America (Vuilleumier 1985, Nores
and Yzurieta 1980, Fjeldsa 1988, Fjeldsa and Krabbe 1990,
pers. obs. J.A.K. and J.A.H., F. Olmos pers. comm. 1990,
R. Schlatter pers. comm. 1990). It occurs from sea level up
to about 3000 m, though a specimen in the US National
Museum was collected at 4200 m, and it occurs in the Puna
Zone at 5000m in the Andes (Fjeldsa and Krabbe 1990,
pers. obs. J.A.K.).
This species feeds primarily in relatively dry, short grass,
such as on relatively dry savannas, agricultural fields, pas-
ture land, and scrub lands in Patagonia and other parts of
southern South America, and is the most terrestrial of the
Neotropical ibises (Nores and Yzurieta 1980, Kushlan et al.
1985, pers. obs. J.A.H. and J.A.K.). It uses very dry ground
during the dry season and after fires, but also makes use of a
variety of damp habitats when available, including edges of
lakes, ponds and rivers, damp meadows, and openings in
forests. Montane populations occur in boggy glades fringed
with forests of the valdivian (rain forest) zone and in grass-
lands of the high altitude Puna and paramo zones. J. Fjeldsa
(pers. comm. 1990) points out that the species is highly
localized in the montane grasslands. Along the coast, it
feeds in shore meadows. F. Olmos (pers. comm. 1990) did
not see them feeding in or around ponds or lagoons.
The birds feed alone, in pairs or in variably sized, but
generally small, loose flocks of up to 50 individuals, which
travel together between feeding sites. They are often
reported in small flocks of up to 6-8 birds in the llanos. We
have seen them feed there primarily in pairs and in the

scrublands of Patagonia we found them feeding singly and
in pairs (pers. obs. J.A.K. and J.A.H.). In the pantanal, F.
Olmos (pers. comm.) found them feeding in pairs and in
groups of up to 4. The montane forms are almost always
solitary or in pairs. It is likely that they primarily remain in
family groups. Brooks (1991) estimated their territories to
average 0.65-1.15 km2 in the Paraguayan Ghaco.
Feeding is by Probing in soft soil and Pecking at the
surface of hard soil. (Sick 1985, R. Schlatter pers. comm.
1990, J. Fjeldsa pers. comm. 1990). In Probing, the birds
can drill deeply into loose soil. On hard ground, they probe
into bunch grass tussocks.
The species consumes a variety of prey found in and on
the soil (Darwin 1962:166, Hudson 1920, MacDonagh 1942,
P.S. Humphrey et al. 1970:103, San Martin 1959, Sick 1985,
Robert Storer pers. comm. 1990, R. Schlatter pers. comm.
1990). The diet is primarily soil invertebrates, including
grasshoppers, crickets, beetles, caterpillars, insect larvae,
spiders, centipedes, scorpions, earthworms and also some
vertebrates, including frogs, toads, tadpoles, salamanders,
lizards, snakes and rats. It swallows all kinds of items, such
as stones, seeds, grass pieces and bones, perhaps as a grit
(R. Schlatter pers. comm. 1990). On Tierra del Fuego a
rodent tooth and shell fragments of a goose egg have been
reported from a large chick (P.S. Humphrey et al.
1970:103). Gantz found that they consume 590 insect larvae
per day (R. Schlatter pers. comm. 1990).
Their flights to roost have been often remarked upon.
They are heard before being seen and repeatedly take the
same flight paths at night to their roosts in trees. These are
often the largest trees around. Sick (1985) reported that they
roosted in parana pines (Araucaria angustifolia) and in the
Amazon estuary in kapoks (Bombax munguba). We have seen
them roost in palms (pers. obs. J.A.K.).
BREEDING
Buffnecked Ibises nest solitarily or in loose colonies of 10-30
pairs situated in a wide variety of locations (A.W. Johnson
1965, Sick 1985, P.S. Humphrey et al. 1970:102). They are
sometimes found in mixed colonies of night herons, cormor-
ants and other wading birds (J. Fjeldsa pers. comm. 1990),
but more usually alone. In the high Andes and on rocky
islands, they nest on the ground and on cliffs. At lower
forested elevations, they nest in trees or rocks scattered on
the savanna or in woods. They have also been reported as
nesting in marshes, on abandoned dredges, on poles, in
reeds in a lake-edge marsh and on the ground along a
stream.
The nesting season varies but usually corresponds to
rainy periods or, in the mountains and in subantarctic
areas, to warm periods (C.S. Luthin pers. comm. 1990). In
the llanos, nests may be found year-round though they are
most frequent in March and October, at the end of the
drying season and in the wet season (Hilty and Brown
1986:72, C.S. Luthin pers. comm. 1990, E. Aguilera pers.
comm. 1990). B.T. Thomas (pers. comm. 1990) observed
birds in the llanos copulating in November.
The southern populations nest in September-December,
Buflfnecked Ibis 187
including those in extreme southern Chile and Argentina
(A.W.Johnson 1965, P.S. Humphrey et al. 1970:103, Nores
and Yzurieta 1980:27, Fjeldsa and Krabbe 1990). In Peru,
they nest in June. In Tierra del Fuego, they arrive at the
time of the first break up of winter ice, nesting in November
to February.
Almost no information is available on courtship and nest-
ing behaviour. B.T. Thomas (pers. comm. 1990) reported
observing allopreening and copulation involving a pair of
birds that were otherwise occupied by feeding in a dry field.
The male mounted the female, they snapped their bills
together, then the male nibbled at the female's neck
feathers. Afterwards, the female walked away shaking her
feathers, and both resumed feeding. Clearly, much more
needs to be known about courtship in this species.
The nest is a large, bulky, loose structure of sticks, with
grass or rush stems intermeshed. Nests are usually placed
high in trees, 3 to 10m above the ground. The nesting site
may be reused in subsequent years, if it is not disturbed
(Vigil 1973:50).
The usual clutch is 2-3 eggs, which are white to greenish
white with some light brown speckling. The incubation
period is 28 days, and in captivity the young are capable of
fledging at 8 weeks (Blaauw 1917).
TAXONOMY
The Buffnecked Ibis is a highly variable species with respect
to size and plumage coloration. As a result the relationship
among the several populations has been much debated, with
several different species and subspecies having been pro-
posed at various times (Salvador! 1900, Todd 1948).
Although some populations have distinctly different colo-
ration, their ranges and intergradations are not well under-
stood. It is likely that most variation is clinal, and it is
unclear how or whether the species might best be sub-
divided taxonomically.
CONSERVATION
The Buffnecked Ibis has long been known, perhaps first
being noted by Cook in Tierra del Fuego, who reported a
species of curlew as big as a heron (Crawshaw 1907). The
species is widespread and abundant over a large part of its
range, especially in the humid pastures and grasslands of
the llanos in the north and pantanal in the south. Overall, it
has adapted well to agricultural practices in the grasslands,
making heavy use of pastures and fields; as a result popu-
lations have increased in southern Chile in the past 40 years
(R. Schlatter pers. comm. 1990).
However, montane populations from Ecuador to Bolivia
are highly restricted. Populations in coastal Peru and north-
ern Chile have decreased, and the bird can only be seen in a
few places (Hughes 1970, Koepcke 1970, Gantz in R.
Schlatter pers. comm. 1990, J. Fjeldsa pers. comm. 1990).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 311.
 Green Ibis
Mesembrinibis cayennensis (Gmelin)

Tantalus cayennensis Gmelin, 1789, Syst. Nat., 1, p. 652; based on Courly verd, de Cayenne of
Daubenton, 1765-81, Planches Enlum., p. 820: Cayenne, French Guiana

Other names:

Cayenne Ibis, River Ibis, Bush Ibis, Curry-curry, Bush Curi-curi, Bronze Ibis (English); Corocoro
Negro, Corocoro de Monte, Cocoquam, Zanurita, Garza Negra, Cocli Negro, Bandurria Habladora,
Caracolero Verde-negro, Cuervillo Grande (Spanish); Korokoro (Dutch); Chapeu-velho, Corocoro,
Tapicuru, Carauna (Portuguese)

IDENTIFICATION
The Green Ibis is a relatively small ibis (45-60 cm long), somewhat stocky, short-legged and long-
billed, with a glossy dark plumage.
The adult's base colour is black, which is how it appears in poor light. However, its entire back is
highlighted by a bronzy green sheen. The forehead is greyish. The green plume-like feathers on the back
of the neck are a distinctively iridescent emerald, but the bushy nuchal crest formed by these feathers is
not terribly conspicuous. The tail and flight feathers are deep blue-black. The ventral plumage is duller
than the dorsal.
The bare face and orbital areas are grey, the bill is olive, and the legs and feet are pea green (S. Olson
pers. comm. 1990). The iris is pale brown to black (Luthin 1983a, S. Olson pers. comm. 1990).
Storrs Olson (pers. comm. 1990) found a bird in breeding condition to have intensified soft-part
colour. The legs and feet were a sparkling metallic green, whereas the throat skin was a violet indigo.
When the gular pouch is distended during courtship, this shows its bright blue cheek-line.
The sexes are alike. Museum specimens labelled as to sex overlap in size, although the largest birds
tend to be males. 2 females collected by S. Olson (pers. comm. 1990) weighed 700 and 740 g whereas 2
males weighed 745 and 800 g. A bird whose sex was not determined weighed 890 g. Bill-length overlaps
similarly. Additional information is needed to determine whether dimorphism occurs in this species.
Nestlings are dark grey, with a distinctive circumorbital white line (C.A. Wood 1923). By fledging,
the juveniles are very much like adults. Luthin (1983a) reported that they have adult soft-part colours, a
green sheen on the feathers, and even a slight nuchal crest. The white eye-line is often apparent in
young for some time after fledging (B.T. Thomas pers. comm. 1990), although it eventually fades with
age.
The Green Ibis is a noisy bird, well known to lockl inhabitants by its dawn and dusk calling, which
gives it and other ibises their most common Spanish name, Corocoro. At night, in the morning and
other times at roost, it is especially noisy. When perched in trees, it often gives a long, low, throaty,
mammal-like growl on an even pitch, repeated for minutes on end (S. Olson pers. comm. 1991).
The primary call of this species is a growling, rattling, rolling 'Kro-kro-kro -kro' or 'Clu-clu-clu-clu'.
When feeding it is quieter, but clucks a throaty, watery 'Koke' or 'Uk' (Slud 1964). Green Ibises often
fly from feeding site to feeding site in small groups in a jerky, heavy, stiff-winged flight. Its flight call has
been variously described, as a rattling 'Klaw-klaw-klaw', an accented 'Wok-wok-wok-wok', and a series
of 3-6 loud 'Ga-lups' followed by 4^9 'Brups' (the latter described by Luthin 1983a).
The Green Ibis is generally recognizable by its size and coloration. It is stockier, shorter and has a
thicker bill than similar species. The Glossy and Whitefaced Ibises have maroon iridescence rather
190 Green Ibis
than green and relatively longer legs. The Barefaced Ibis is
smaller with pink rather than green soft parts. The Plum-
beous Ibis is grey.
DISTRIBUTION AND POPULATION
The Green Ibis is a forest bird, primarily of the wooded
basins of the Amazon and Orinoco rivers of South America,
a habitat preference reflected in several of its older common
names, such as River Ibis and Bush Ibis. Its range extends
northward into Central America. In the north it ranges from
the Caribbean coast of Costa Rica to Limon, through
Panama, northern Colombia to the Venezuelan interior
(Orinoco and Apure Rivers) as well as coastal Falcon, the
Guianas, west again to east of the Andes in Brazil, Bolivia
east to Beni, Paraguay, and northern Argentina in Misiones
(Olrog 1963:88, D.A. Scott and M. Carbonell 1986).
Seasonal population shifts seem to occur in some areas. In
the pantanal, individual Green Ibises are reported to occur
in May-November, and in the llanos from November to
August (B.T. Thomas 1979a, R. Cintra pers. comm.). Indi-
viduals are frequently found outside the nesting areas. Dis-
persing birds are found as far northward as Sarapique,
Costa Rica and Rio Plantano, Honduras (American Orni-
thologists* Union 1983) and stragglers in Rio Grande do
Sul in January (Sick 1985).
Nothing is known about population sizes and the birds
are well dispersed in suitable habitat. However, they are
locally common, and overall population levels are probably
as high as could be expected for such a bird,
ECOLOGY
The Green Ibis is a bird of the water's edge and shallow
water. However, it is found locally in a variety of habitats,
including wooded swamps, forest pools, small marshes, hea-
vily timbered gallery forests along rivers, on forested
streams having muddy banks and along roadside ditches. It
also occurs in secondary growth, such as abandoned coffee
plantations (Haverschmidt 1968) and, as we have seen, in
more open pasture as well (pers. obs. J.A.K.). It feeds in
muddy areas particularly, on the edge of river banks and
forested pools, where its dark coloration renders it well
camouflaged.
Generally, this is a rather solitary bird. In a study of its
feeding habits in the llanos, one of us found it feeding
solitarily, in pairs, or in small loose flocks of up to 6 or 8
birds (pers. obs. J.A.K.). Gonzalo Morales (pers. comm.
1990) observed groups of up to 30 feeding in the Venezuelan
llanos. Betsy Thomas (pers. comm. 1990) has also seen
flocks in the same region, reporting one of 26 birds. When it
feeds near other species of ibis—though this is infrequent—
it is usually near the edge of the group (pers. obs. J.A.K.).
Green Ibises appear to use single feeding areas and roost
sites for much of the year. They seem to establish territories
that they appear to claim by calling. Luthin (1983a) found
only solitary birds or pairs during the nesting season; but,
after nesting, the larger groups of 2-6 typically seen are
probably families. Often the individuals of a probable pair
are well separated when actually feeding, but they and the
larger family group roost together at night.
Upon disturbance, the Green Ibis turns its head jerkily
while nodding and making its clucking sound (Slud 1964).
It then walks or flies to nearby trees where it calls in its
distinctive manner, often hidden from view.
Green Ibises forage for long periods throughout the day,
and spend the remainder perched in nearby trees. They feed
by walking relatively quickly, Picking at the surrace and
Probing rather deeply in soft mud (pers. obs. J.A.K.). They
also feed while rapidly opening and closing their wings
(Ogden and Thomas 1985a).
A muddy feeding spot is characteristically chosen, and
birds are often seen with bills partially mud-covered. We
have observed them feeding slowly by a sequence of
Standing-Probing-Walking-Probing in a thorough and de-
liberate manner, completely working over a patch of mud, a
small pool or a stream edge, spending many minutes or an
hour in a single area (pers. obs. J.A.K.). This deliberate
foraging tactic has been well known for many years (R.H.
Schomburgk 1848:503). Luthin (1983a) reported numerous
probes around soft earthworm holes in the forest.
Green Ibises are documented to eat aquatic insects
(Orthoptera, Homoptera, water bugs and water beetles),
worms, molluscs, including snails, small fishes and frogs
(Sick 1985, Ogden and Thomas 1985a,b, G. Morales pers.
comm. 1990, pers. obs. J.A.K.). Sick (1985) has reported
two instances of its eating plants. He found large quantities

of fibrous vegetable matter in two specimens taken at Lin-
hares, Esprito Santo, Brazil (H. Sick pers. comm. 1990).
Undoubtedly, the overall diet is more varied than what has
so far been recorded.
BREEDING
Green Ibises nest solitarily and inconspicuously high in
trees and thickets, usually in gallery forests along rivers.
They appear to nest within their long-held activity areas.
Birds in breeding condition have been collected in
Panama and Colombia in February-April (S. Olson pers.
comm. 1990, US National Museum specimens). They have
been reported as nesting in April in Colombia, September in
Surinam, and June-August in Venezuela (Haverschmidt
1968, B.T. Thomas 1979a, 1990, Hilty and Brown 1986:73).
In the llanos, courtship and nesting occurs during the rainy
season, typically beginning 2 months after the onset of the
rains (Luthin 1983a). Its nesting season is shorter and more
consistent than that of the Barefaced Ibis, which nests in the
same area.
Luthin (1983a) was able to study some of the displays of
this species, providing most of what is known and described
below. It is likely that birds remain paired year-round, so
some courtship displays may be truncated, and the court-
ship period is rather extended. As the breeding season
approaches, display sites are selected at the top of tall trees,
probably within the pre-existing home range.
During courtship, Green Ibises alight together on a dis-
play tree, standing no more than 2 m apart. Courtship is
initiated by one bird performing a Head Shaking display,
and the second bird responding with its own Head Shaking.
The pairs then may Duet, the call consisting of a soft, deep
'Brrr*. They hold their necks near vertical and elevate their
head feathers. At the end of the duet, both birds shake their
heads sideways and bump bills. One or both birds may flare
their gular pouch, showing the bright blue line along the
cheek.
After a pause some birds then begin Stick Shaking or
bouts of allopreening directed to the head and neck.
Occasionally, Display Preening occurs, in which the birds
raise the feathers on their necks and back and slightly droop
their wings. Display Preening and Stick Shaking are less
common in the Green Ibis than in other, similar species
(Luthin 1983a).
Copulation displays are, similarly, less developed than
some of the other Neotropical ibises. The male Green Ibis
stands close to the female, sweeps his bill back and forth in
front of her sometimes with his head over hers. The male
tries to grab the female's bill while mounting her, and may
keep hold during copulation.
The pair constructs a rather flimsy, flat structure made
loosely of sticks, lined with finer material, according to C.A.
Wood (1923), although Luthin (1983a) failed to find such
finer material in the nests he examined.
The birds are secretive in their nesting, entering and
leaving the nest-site quietly. Luthin (1983a) found that
Green Ibis 191
activity at the nest was limited and vocalizations consisted
only of a quiet gurgle during exchanges at the nest.
Eggs are a dark olive green, without markings or gloss. 2
eggs from Surinam measured 54.9 X 38.8 and 55 X 37.9 mm
and weighed 46 and 43 g (Haverschmidt 1968). Clutch size,
when recorded, has been 2-4. Hatching is asynchronous.
Luthin (1983a) found that only 2 young survive to fledging.
Hatchlings have distinct facial markings that disappear in
2 or more weeks (Luthin 1983a). Like the adults, young also
are quiet at the nest, begging with a light buzzing call (C.S.
Luthin pers. comm. 1990). After 2-3 weeks, the adults re-
turn to the nest-site only to feed the young, which continue
to remain relatively inactive.
Young in the nest crouch down when approached by
humans (C.S. Luthin pers. comm. 1990). The secretive
behaviours of the young and adults at the nest suggest that
predation risks may be high. Luthin (1983a) relays a report
of capuchin monkeys (Cebus nigrivittatus) eating young.
Other predators are undoubtedly a threat also.
The young fledge at 23-27 days. They remain near the
nest for a few days and then move with the adults to their
feeding sites, thereby forming the family groups so apparent
at the end of the rainy season and thereafter.
TAXONOMY
Like most ibises, the taxonomy of the Green Ibis has not
been thoroughly studied. Based on behavioural information,
Luthin (1983a) has suggested that Mesembrinibis might be
related to the solitary Old World ibises of the genus Bostry-
chia, such as the Spotbreasted, Olive and Hadada ibises.
Both groups are secretive, solitary, forest-dwelling species,
loudly calling at dawn and dusk, having dark plumage and
nuchal crests, and building loosely constructed nests high in
trees. Behavioural patterns held in common include mutual
Head Shaking while bumping bills and elevating the
feathers, bill grasping during copulation, and Mutual
Preening. However, these behaviours are not unique, and
additional behavioural and morphological study is called
for.
CONSERVATION
Little is known about the biology, detailed distribution and
population sizes of this species. Such information is needed
to understand its ecological requirements and its conser-
vation needs. However, the Green Ibis appears to be widely
dispersed within its range, and is, overall, probably reason-
ably common.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 312.
 Hadada Ibis
Bostrychia hagedash (Latham)

Subspecies:

Bostrychia hagedash hagedash (Latham)

Tantalus hagedash Latham, 1790, Index Ornith., p. 709: Cape of Good Hope

Bostrychia hagedash brevirostris (Reichenow)

Theristicus brevirostris Reichenow, 1907, Ornith. Monatsber., 15, p. 147: Alen, Southern Cameroon
(=Equatorial Guinea)

Bostrychia hagedash nilotica Neumann

Bostrychia hagedash nilotica Neumann, 1909, Ornis, 13, p. 193: Kimo, NW of Addis Ababa, Ethiopia

Other names:
Hadada, Hadedah (English); in-Kankane, 1-Han, Ing'ang'ane (South Africa); L'ibis Hagedash
(French)

IDENTIFICATION
The Hadada Ibis is a short-legged, medium-sized (610mm long), dull brown bird, with a somewhat
greyer head and neck.
Close up, the basically brown coloration of the mantle and back of the adult has an iridescent green to
bronze cast. The head is greyer than the body, with a prominent huffish white stripe running across the
cheek. The neck is a brownish or smoky grey. The colour of the primaries, secondaries and major
coverts vary somewhat according to the light, from greenish brown to pale red and purple, forming
coloured patches on the wing. The tail is dark blue, with the rump and tail coverts green.
The bill is dark horn with a strong red marking at the base of the upper mandible. The featherless
lores and base of the bill are black, and eye is dark brown ringed by a prominent white ring. The legs
and feet are dark horn-brown. During courtship, the red marking on the bill brightens to a crimson, and
the bill itself darkens. The legs and feet turn darker, and a pale reddish flush appears on the toes.
The nestlings are pink with brown down, black bill, and pink legs and feet. Immature birds are
similar to adults but are greyer and have less iridescence on the back and wings.
The best-known characteristic of this ibis is its jerky irregular flight and its loud echoing call, 'Ha-da-
da', from which it gets its common English name. This call is given by both male and female and is
particularly heard when flying to and fro between the roost and feeding ground. It is also heard quite
often during the day, as birds move between feeding grounds. It feeds silently and allows quite close
approach where not persecuted.
The main differences between the three subspecies are size and some aspects of coloration (Amadon
1953). The hagedash subspecies is relatively short-billed and paler than the others. The brevirostris
subspecies is a darker brown colour with its glossy wing patch much greener and more purple,
according to the light. The bill of nilotica is noticeably longer, and some specimens are strikingly larger.
(Comparative measurements are: hagedash—bill 11.7-15.3, wing 33.4-37.Ocm; brevirostris—bill 12.6-
16.3, wing 33.0-38.3; nilotica—H\\ 15.2-17.4, wing 33.5-39.2—Brown et al. 1982.)
This species is easily confused with others in the genus Bostrychia, as noted in those accounts. Voice
differences are important in identification. The Hadada has a very noisy, harsh call with three distinct
syllables. It has been compared to the wail of a child in distress and the hideous mocking laughter of a
witch (F J.Jackson 1938). The three part call contrasts with the two syllable call of the Olive Ibis. The
194 Hadadalbis
Hadada's call also contrasts with the more raucous call of
the Spotbreasted Ibis. The other species of Bostrychia also
have a very prominent nuchal crest, which is minimal in the
Hadada Ibis.
DISTRIBUTION AND POPULATION
This ibis is a widespread species throughout the African
continent south of the Sahara desert (Mackworth-Praed and
Grant 1962, Snow 1978, Brown et al. 1982:195-196); hage-
dash is found in Zimbabwe, southern Mozambique, and
eastern South Africa. North of the Zambezi Valley, hagedash
is replaced by brevirostris, which is found throughout central
and east Africa, northwards through Kenya and Somalia
and westward to Gambia and Sierra Leone, and along the
Congo River.
The nilotica subspecies occurs from Uganda and western
Tanzania to Zaire, Sudan and Ethiopia (Pitman 1928,
193 Ib). Near Mount Kenya, it is a lowland species and is
replaced at higher altitudes by the shyer, rarer Olive Ibis.
In southern Africa, the Hadada has in recent decades
expanded its range from the southeastern coastal belt to the
two higher inland plateaus, as well as westward along the
coast (Skead 1966). This expansion has been in response to
the construction of dams and reservoirs (Snow 1978).
Little is known about the population sizes of this species,
despite (or because of) its wide range. Bannerman (1953)
found it to be a common bird in certain areas of west Africa
but rare in some coastal provinces, particularly in Nigeria.
A colony in Cape Province, South Africa, was recorded as
numbering 70 birds (Raseroka 1975a).
ECOLOGY
The Hadada Ibis inhabits damp ground along rivers and
streams, especially preferring open woodland, savanna and
marshlands near trees. It is attracted to lakes and man-
made irrigation, avoiding the desert regions of north and
southwest Africa as well as thick forest areas. It is found
close to farms, cultivated farmland and lawns. On Mount
Kenya and other mountain areas, it is not found above
1800m.
Hadada Ibises are, in the main, sedentary birds feeding in
pairs, small groups or in some cases parties of up to 50 birds,
sometimes in association with other birds, especially Cattle
Egrets (Bubuleus ibis). We have also seen single birds quietly
feeding in shallow pools in the company of Goliath Herons
(Ardea goliath), Wood Sandpipers (Tringa glareola) and
Greenshanks (Tringa nebularia).
Flocks vary in size during the day, birds breaking away
periodically to feed elsewhere. In a tree-lined marshy pool in
the Masai Mara in Kenya, we have watched Hadada Ibis
arrive in small parties, noisily flying in at dawn, but then
settling to feed quietly throughout the day, until they move
off to feed around village huts on their way to their roost.
Between feeding sites, they fly haphazardly with jerky
flaps.
Preferring open ground, they feed on both dry and damp
areas of short grass and cultivated fields and even gardens,
usually by Probing in soft soil. On hotel lawns along the
shoreline of Lake Naivasha in Kenya, we have seen them
Walking Slowly in small parties in company with Sacred
Ibis, Probing the grass and flower beds and vegetable
patches. They particularly favour the base of trees where the
soil is exposed.
The availability of soft soil may be critical to the species.
Ossowski (1952) suggested that heavy mortality during
severe droughts was due to the hardness of the soil. The
birds may also be surface feeders under some conditions
(Raseroka 1975b), but this needs additional study.
Hadada Ibises are attracted to wattle plantations in
Natal, South Africa, and are thought thus to act as effective
pest controllers (Ossowski 1952). This work also underlined
the value of the Hadada in keeping golf courses free of the
larvae of chaffer beetles (Ossowski 1952).
The Probing extracts a variety of invertebrates, including
earthworms, beetles, Lepidoptera, fly larvae, millipedes,
water bugs and crustaceans. Raseroka (1975b) found that
96% of the animals ingested were insects, and 60% of the
diet was made up of animals obtainable from the soil sur-
face. They have not been recorded as taking large prey. Bell-
Cross (1974), in his observations offish eating birds, never
found this species to eat fish.
Inedible items, such as stones and glass, are often found
to be taken by Hadada Ibises (Skead 1951, Ossowski 1952).
Skead (1951) concluded that they were attracted by bright
objects on the ground.
As evening approaches, birds straggle to roosts, some
pairs roosting apart from the main flock. At the roosts,
the birds will call loudly to each other and will be answered
by birds at other nearby roosts (Manry in Brown et al.
1982: 195-196, pers. obs. J.A.H.). These large nocturnal
roosts are of 100 or more birds, usually away from other
species. They are often used year-round, and year after
year.

BREEDING
It is likely that some birds maintain a pair-bond through the
year, as we have seen interactions outside the nesting season
that suggest this. Commonly, a female, with wings half
open, approaches a male and touches her bill to his. After
this brief mutual bill touching the female will resume feed-
ing elsewhere amongst the party. When disturbed, the two
birds will separate from the other birds and fly up to the
branch of a tree standing close together.
Hadada Ibises nest solitarily in trees or bushes, often over
or near water (Raseroka 1975a). In South Africa, nests have
been seen in trees some distance from water, and even on a
telegraph pole (Line 1941, Raseroka 1975a, Uys and Broek-
heysen 1966).
Time of nesting varies geographically. Raseroka (1975a)
showed that nesting ranged from February in Zambia to
October-November in southern Africa (also observed by
Ossowski 1952). In Eastern Cape Province, eggs were laid
from the beginning of November to mid-January. Nesting is
recorded in April in Uganda (Pitman 1931b).
As the breeding season approaches, paired birds will
separate from the feeding parties, which gradually break up
as the nesting season commences. It is likely that estab-
lished pairs have a lower intensity of courtship interaction
than do those nesting for the first time. We saw one such
incident at the commencement of the rains in Kenya, and it
was noticeable that red parts of the upper mandible were
brighter on the interacting pair than others of the feeding
party.
In a party of 8 feeding birds, one bird picked up single
sticks, grass and straw at intervals, and paraded through the
group and then away from it. It then crouched amongst the
other birds, vibrating its half-open bill. One of the birds
then came from the group and stroked the quivering bill of
the displaying bird with its own closed bill in a 'crossed
sword' action. The first bird then touched the prominent
white stripe on the second bird and the interplay ceased. We
saw this behaviour by the two birds about eight times in
over 15 min.
Skead (1951) noted that the billing behaviour of pairs
included rattling of bills, up and down, and side to side,
while nodding the head. This took place at roosts, but more
intensively at nesting sites, where sticks were offered by each
bird to the other, followed by neck intertwining, mutual
preening, head shaking and jibbering of the lower mandible.
W. Wennrich (1983) noted that this behaviour preceded
copulation and was followed by mutual head raising and
lowering in unison. Stick presentation continues into the
nesting period, and during incubation the arriving bird will
offer a stick to the sitting bird, which will then leave the nest
without further display between the pair.
Large sticks form the base of the nest and these are lined
with small sticks and then with grasses and lichen. Nests
can be completed within 14 days but can take up to a month
or longer, with periods of suspended activity (Raseroka
1975a, Brown et al. 1982:195-196). Sticks are both removed
and added as building goes on. Nests in Combretum caffrum
trees averaged 17 m apart and varied from 1.2 to 7.0 m high
Hadadalbis 195
above the water or ground (Raseroka 1975a). Nests are used
for a number of years, and Skead (1951) saw a nest in a
kaffir plum tree used for 6 consecutive nesting seasons,
although there is no certainty that the same pair used the
nest each year. Some birds will take over partly built nests.
2-3 eggs are laid, but sometimes 4, on alternate days.
Eggs are large, often heavier than those of a domestic fowl,
but highly variable even within a single clutch. They are a
buff colour and variably spotted and flecked with light
brown. Often the flecks are overlaid with sepia pigment.
Size varies from 56.9 X 41.5 to 64.0 X 41.0mm, with an
exceptional egg measured at 67.2 X 42.5 (Skead 1951). Pit-
man (1928) noted an average egg size of about 60 X 42 mm.
Incubation takes between 25 and 28 days and is shared by
both parents (Raseroka 1975a). Chicks hatch at long inter-
vals, some taking up to 48 h to leave the egg, and thus the
developmental stage of nestlings vary. Young remain in the
nest for up to 33 days. They are blind for the first 3 days and
have pink skin covered in brown down. Their bill is black
and straight. They utter a purring sound when food beg-
ging, and are fed by regurgitation, which continues after the
young have left the nest. Before fledging, young are left at
the nest, but are brooded at night (Raseroka 1975a). Each
parent takes young to feeding areas, coming together as a
family regularly and roosting together in the vicinity of the
nest.
TAXONOMY
With the merging of the genus Hagedashia into Bostrychia, the
relationships of the Bostrychia ibises seems well settled. The
Hadada Ibis is polytypic, the several subspecies differing
somewhat in size and colour. In fact, another race, B. h.
erlangeri was once recognized, which is now included in
brevirostris:
Related to this diversity, an interesting aspect of the
Hadada Ibis is that the structure of its chromosomes (its
karyotype) differs from that of other ibises, which is surpris-
ingly variable anyway (see Classification). Capanna et al.
(1982) have noted that karyotypic variability is found in
taxa that are characterized by strict territoriality, small
individual populations and gregariousness, traits that en-
courage inbreeding. These results suggest the possibility the
Hadada Ibises have a higher rate of inbreeding than might
otherwise be expected, leading to interesting questions of
breeding biology and conservation.
CONSERVATION
This species has acclimatized well to human habitation
(Wolstenholme 1961), and in fact has been a part of human
culture for centuries (Schiiz 1970a). It is not subject, over
the majority of its range, to persecution. The main danger to
its population stability comes from extended droughts. Con-
versely where additional water tanks and ponds are con-
structed, this species will move in to colonize the newly
irrigated areas where trees are available.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendi"
dix —
on
page 313.
 Wattled Ibis
Bostrychia carunculata (Riippell)

Ibis carunculata Riippell, 1837, Neue Wirbelthiere Fauna Abyssinien, Vogel, p. 49, pi. 19: Taranta
Mts, Ethiopia

IDENTIFICATION
A medium-sized, dark brown, crested bird, the Wattled Ibis is distinguished by the combination of its
dark colour, its crest and its white wing-patch. It has a relatively short bill, a relatively long tail, and a
small wattle hanging from its throat.
The basic colour of the adult is dark brown, darker above than below, with a greenish gloss on the
back. The crown is dusky, and a crest of the same colour extends behind the neck. The wing-feathers are
black and glossy, the primaries having a bluish gloss and the secondaries, a greenish gloss. The upper
wing-coverts are also black, except for the white wing-patch. The brown face is feathered with a dusky
red bill. The red eye is surrounded by a white ring. The wattle is a thin, dusky red pendant, up to
20 mm long, which hangs from the lower throat. The legs and feet are also dusky red.
The bill-length has been given as 11.0-12.7 cm (Friedmann 1930) and as 8.3 cm (de Naurois 1973),
the difference suggesting that there may be a sexual dimorphism in the species. Friedmann also gives
the wing-length as 32.5-38.0 cm and the tarsus as 6.3-6.6 cm; de Naurois gives the wing length as 32.5-
38.0 cm.
During courtship the soft-part colorations of the bill, wattle, legs and feet brighten.
Nestlings are covered in black-brown down. They have bare skin around the eyes; their straight bill is
grey with a black tip (Dorst and Roux 1972).
In flight, like the other Bostrychia ibises, the Wattled Ibis is very noisy. Flying to and from roost, often
in flocks, it gives a cry described as 'Kowrr-kowrr-kowrr'. This is louder than the call of the Hadada
and, in a chorus, can be heard up to 5 miles away. Besides the raucous calls, roosting birds utter a series
of grunts, groans, and squeaks (Urban 1978).
The Wattled Ibis may be confused with other Bostrychia ibises. It is larger than the uncrested Hadada
and is recognized in flight by its white wing patch. At closer range, the red pendant shows clearly. Its
crest is less well developed than those of the Spotbreasted and Olive ibises but more developed than the
uncrested Hadada Ibis.

DISTRIBUTION AND POPULATION

This species is resident all over the Ethiopian plateau of northeastern Africa. It occurs from about
1500 m to the high moorlands of about 4100 m elevation. The population has not been censused, but it
has been common within its restricted range, especially near Addis Ababa and Bale. Flocks of over 100
birds were commonly reported.

ECOLOGY
Difficulty of access to the Ethiopian plateau region has meant that few observations have been made of
the ecology of this species. The birds appear to be sedentary, often roosting near the breeding sites.
They roost in trees singly, in pairs or in small groups, and sometimes in larger groups. They also roost
on rock cliffs some distance from water (Brown et al. 1982:196-197). After dawn they fly to the feeding
198 Wattled Ibis
grounds, where they forage singly or in flocks of up to 100
birds.
The Wattled Ibis feeds in open grasslands, marshes,
alpine woodlands, among crops or in forest glades down to
1500m altitude (Brown et al. 1982: 196-197). Their forag-
ing behaviour is Walking while Probing the ground regu-
larly. They will accompany herds of domestic animals,
turning over dung heaps for beetles. If disturbed, they will
rise as a flock, calling loudly. Their food consists of small
prey including worms, insect larva, other insects, molluscs,
frogs, small reptiles and mammals.
BREEDING
Nests are usually built in small or large colonies on cliff
edges at over 2400 m altitude. Occasionally they nest at
lower altitudes, singly or in small groups of 2 or 3 pairs, in
trees or even on the ledges of buildings (Fry et al. 1985).
Schiiz (1970a) found a nest-building pair near Lake Awasa
at 1700 m, which he considered was probably near the lower
limit of nesting.
The nest is made of branches and large sticks, lined with
softer plant material. It is c. 20-50 cm across. On cliff edges,
the nests may be close enough to touch or may be several
metres apart. The breeding birds often choose to nest on
eastern slopes, probably so as to obtain maximum early
morning sun, because night-time temperatures at the alti-
tudes at which they nest can be very cold (Dorst and Roux
1972).
Males claim and defend territories, and females remain at
the nest site. Laying dates are well dispersed, March-May,
July and December, which include the small rains, the big
rains and the dry season. The eggs were described by de
Naurois (1973) and by Brown et al. (1982: 196-197) as dirty
white, rough-shelled and marked with brown. On the other
hand, Dorst and Roux (1972) described them as being olive
green with brown markings. The eggs are 59-61 X 39-
40cm and weigh c. 50 g. Upon fledging, the young feed
away from the main colony.
TAXONOMY
The Wattled Ibis is the only montane, non-forest species of
Bostrychia. Its habit of nesting on rocky cliffs and ledges is
similar to the Geronticus ibises, and in this it may show
considerable affinity to the latter genus (Fry et al. 1985).
CONSERVATION
Although previously a common species, within its very
restricted range, long periods of recent warfare and drought
raise concerns for this ibis. Little is known about its present
status.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 314.
 Olive Ibis
Bostrychia olivacea (Du Bus de Gisignies)

Ibis olivacea Du Bus de Gisignies, 1838, Bull. Acad. Roy. Sci. Lettres Beaux-Arts Beligique, 4 (1837),
p. 105 and pi: 'Cote de Guinee' (=upper Guinea, as determined by Chapin, 1923, Amer. Mus. Novit.,
no. 84, pp. 1-3)

Subspecies:

Bostrychia olivacea olivacea (Du Bus de Gisignies)

Bostrychia olivacea cupreipennis (Reichenow)
Theristicus cupreipennis Reichenow, 1903, Ornith. Monatsber., 11, p. 134
Bostrychia olivacea rothschildi (Bannerman)
Lampribis rothschildi Bannerman, 1919, Bull. Brit. Ornith. Club. 40, p. 6: Roca Infante D. Henrique,
Prince's or Principe Is.
Bostrychia olivacea bocagei (Ghapin)
Lampribis bocagei Chapin, 1923, Amer. Mus. Novit., no. 84, p. 5, figs 2A, 3A: Rio de Sao Tome,
Sao Tome

Bostrychia olivacea akleyorum (Chapman)

Oreoibis akleyorum Chapman, 1912, Bull. Amer. Mus. Nat. Hist., 31, p. 235, pis 23-24: south slope of
Mt Kenya, altitude 9000 ft

Other names:
African Green Ibis, Green Ibis, Kenya Ibis, Dwarf Ibis and Dwarf Olive Ibis (bocagei) (English);
Gallinhola and Gallinho do Matto (bocagei) (Portuguese); L'ibis Olivatre, Ibis Olive a Crete (French)

IDENTIFICATION
The Olive Ibis is a small to medium-sized, dusky brown ibis with a prominent crest and red face and
bill.
The back is a dusky brown with a green iridescent tinge. It is very dark brown below, which also is
tinged with green. The very obvious head-crest is dark brown with a purplish sheen to the long feathers
that extend well back from the top of the head. Secondaries and wing-coverts show green, bronze and
pinkish lustre; the tail is dark blue.
The bare face and bill are coral red, and the red eye is set off by blue-black lores. The legs and feet are
described as yellowish green to pinkish brown and also as dull dark red (Brown et al. 1982). It seems
certain that the differing descriptions of the soft parts refer to birds in different stages of breeding
coloration. These colour changes need to be better documented.
The subspecies differ in details of coloration and geographical range. The subspecies olivacea has faint
pale striping on the head and neck. There is more green on the neck and body of cupreipennis. The crest of
rothschildi is more purple. The subspecies bocagei is duller plumaged and very much smaller than olivacea.
The subspecies akleyorum is larger than olivacea, and its body is noticeably greener. Other than noted
above, there is a considerable overlap in size among the races. In view of the small number of
specimens, patterns of size variation cannot be considered definitively known.
The very limited data for rothschildi suggest that males are larger than females, but this needs to be
confirmed. In other ways there is no indication that sexes differ.

(7) olivacea (lowland forest); @ cupreipennis (lowland forest);
@ akleyorum (mountain forest); (4) bocagei (Sao Tome Island
only).
The nestlings have dark brown down. Immature birds
lack the long crest and are duller coloured than the adults.
The voice of the Olive Ibis is a strong resonant double
squawk, rendered *Gar-wa, gar-wa*, and also a goose-like
call heard at a long distance (Brown et al. 1982). When
annoyed it will give call 'Gar' at an intruder (Meinertzha-
gen 1937).
This ibis has a straggling flight and travels high over the
canopy at dawn and dusk calling loudly as it flies.
The Olive Ibis looks very similar to the closely related
Hadada and Spotbreasted ibises. Although the Spot-
breasted Ibis has its neck and underparts well spotted and
also has a green spot in front of the eye and a green streak
crossing the face in line with the bill, A. Brosset (pers.
comm. 1990) has stressed to us that the Olive and Spot-
breasted Ibises are practically impossible to distinguish by
plumage and size in the field. Similarly, many observers
have confused the Olive Ibis with the more familiar
Hadada, especially when the Olive Ibis is in deep or mottled
shade, where little of its iridescent colouring is visible. It
appears to fly more heavily than the Hadada Ibis and
appears to have a thicker neck (Meinertzhagen 1937).
In many cases, voice is the best character for separating
these species. The call of the Spotbreasted Ibis lacks the
honking quality of the Olive Ibis; and, as Chapin (1932a)
pointed out, the Olive Ibis does not stress the second note as
does the Spotbreasted Ibis. The Hadada has three distinct
notes instead of the two of the Olive Ibis. (Tapes of calls are
listed by or available from Chappuis 1980, The British
Library of Wildlife Sounds, and Keith and Gunn 1971.)
Olive Ibis 201
DISTRIBUTION AND POPULATION
This widespread sedentary species has seldom been
reported in the wild, especially in recent years, so very little
is known about the details of its distribution and nothing
about its population levels. It may be more common than
we can document because its habitat is so inaccessible and
because it is so difficult to identify with certainty in the field.
The subspecies olivacea is found in West Africa in Sierra
Leone and Liberia; cupreipennis is also found in West Africa,
in Cameroon, Gabon, Congo and Zaire. The subspecies
akleyorum is found in the mountains of Kenya and Tanzania,
particularly Mount Kenya and the Aberdares, the Usam-
bara Hills, and probably Mount Meru (F J. Jackson 1938,
Snow 1978). Collected only from the Gulf of Guinea Island
of Principe (Prince's), rothschildi is now thought to be ex-
tinct.
The other island subspecies, bocagei, is found only on the
nearby island of Sao Tome. In 1988, John Burlison and
Peter Jones, along with Ana-Maria de Sa Almeida and
Claudio Menezes do Espirito Santo, visited southwestern
Sao Tome and received credible accounts of the survival of
this race (World Birdwatch 1988). In June, 1989, Jan Haft
and Jacob Fahr reported seeing birds of this species along
the lo Grade River, about 4 kilometres north of Ribeira
Peixe (World Birdwatch 1989). The reports of these birds,
the first since a specimen was collected in 1928, are one of
the more exciting discoveries in ibis conservation of the past
decade.
ECOLOGY
The Olive Ibis is a solitary species of deep forest, but has a
rather intriguing habitat distribution over its range (Snow
1978). It is widespread in the lowland forests of West Africa
but is absent from the Cameroon highland forest and from
the mountains forming the eastern edge of the Congo basin.
Yet it is a montane bird in Kenya and Tanzania, where it
occurs upwards from about 1800 metres. In West Africa it
occurs with the Spotbreasted Ibis, but in East Africa it
appears to be replaced at low altitude by the Hadada Ibis
(Snow 1978).
Certainly akleyorum is more often seen than the lowland
races to the west. We found akleyorum near streams in the
montane forests of Kenya. Vincent C. Fayad has reported
seeing it more often on the Sirimon route on Mount Kenya
than on the Naro Moru route. In September 1988, he
encountered 10-12 birds at an altitude of 3000 m near the
Sirimon road head (V.C. Fayad pers. comm. 1989). I.S.C.
Parker (pers. comm. 1990) also reports that this species is
found in some numbers in the Hegenia abyssinica forest of
some 100-200 acres, surrounded by moorland, between
2700 and 4000 m in the Aberdare Mountains.
We have searched, unsuccessfully, for olivacea in the forest
of West Africa, but have only succeeded in attracting very
large numbers of leeches which fall from the trees in large
numbers! However the fast running streams strewn with
boulders, which we found there, mirrored the similar East
African streams where we have observed akleyorum. The
other western subspecies, cupreipennis^ occupies a similar
lowland forest habitat.
202 Olive Ibis
On Sao Tome, Burlison and Jones (pers. comm. 1990)
reported that the endemic bocagei occurs in the remote,
forested regions at high altitude. They reported that local
people had never seen it in tall forest trees grown on long-
abandoned oil palm plantations but only in the 'abo', or
untouched primary forest. In June, 1989, 5 individuals were
observed on lava gravel along the Rio Grande River (World
Birdwatch 1989).
In its forest habitat, the Olive Ibis feeds along the moun-
tain streams and riverlets, although Bannerman (1953)
encountered it in the vicinity of larger rivers such as St.
Paul's River in Liberia and the Gamma River in Gabon.
I.S.C. Parker (pers. comm. 1990) suggests that the Hegenia
forests of Kenya contain a number of birds, and indicates
that they prefer to forage on the forest floor where mature
trees have little or no undergrowth.
The species is described by Bannerman as having regular
habits, feeding in marsh, swamps and backwaters and
returning to high trees, such as a silk-cotton, to spend the
night. It prefers to roost in such very large trees, with dead
tops (Brown et al. 1982). Bannerman (1953) stated that
when disturbed it takes refuge in thickly leaved trees. It
travels high above the canopy at dusk and dawn calling
loudly. It is silent throughout the day when feeding, and so
is difficult to locate.
Nothing is reported regarding feeding techniques of the
Olive Ibis. It appears to feed in pairs or small groups (W.L.
Sclater and R.E. Moreau 1933a). Its food consists of insects
such as beetles, larvae, grubs, snails and worms. A male
specimen from East Africa contained beetles, and a female,
green leaves, beetles and segments of Myriopods (FJ. Jack-
son 1938). It is reported by Brown et al. (1982) to eat snakes
and some vegetable matter from the forest floor.
BREEDING
The earliest, and one of the few, reports of breeding is the
account of Mr and Mrs Carl Akeley who found a nest of the
East African race in 1910 on the southern slopes of Mount
Kenya in dense forest at about 3000 m elevation (Chapin
1923, F.J. Jackson 1938). It was seen in a bamboo forest
near their hunting camp. The nest was loosely constructed
of dead branches on the limb of a small tree about 8 m from
the ground. The nest contained 3 young at the time, and a
fragment of an eggshell which was pea-green and more or
less stained with cinnamon rufous and irregular blotches of
chestnut. It was much less heavily marked than the egg of
the Hadada Ibis (F.M. Chapman 1903).
No further nests were discovered until 1959, when A.
Dyer found one at the headwaters of the Pesi River in the
Aberdares (Kenya) in a mixed Juniperus-Podocarpus forest
(I.S.C. Parker 1982). This section of the Aberdares is rela-
tively dry forest, being in the range's rain shadow, and the
Pesi is a very small stream at this point, no more than 2 m
across. This nest was in the fork of a slender branch over a
pool about 2 m below a small waterfall. The structure was
flimsily made, of large twigs between 30 and 40 cm across.
Both birds were observed building, which is accomplished
silently.
During a visit in 1980, we discussed the matter with
ornithologists in Kenya, remarking on the sparsity of sight-
ings of this species. The next year, Mr I.S.C. Parker was
able to find a nest again in the Aberdares (I.S.C. Parker
1982). The nest was observed in July and September at a
small stream near the Thiririka river, on the southeastern
slopes of the Aberdares in mixed Podocarpus and Ocotea
forest. This nest was also over water, precariously placed
about 4 m high on a very slender branch 2 cm thick and
partially supported by several even smaller branchlets and a
vine. It was about 30-40 cm in diameter, and made of large
twigs such that it could be seen through from the ground. In
late July it was empty (although what Parker thought to be
a leaf when viewed through the nest bottom could have been
an egg), but by the first week in September contained 2 half-
grown nestlings (I.S.C. Parker 1982). When visited by Ian
Parker and Peter Davey later in the month, the nest had
largely fallen down. Since that date, no further observations
have been made due to heavy mists and rainfall at that time
of year (P. Davey pers. comm.). No other nesting records
are known, and there remains no record of nesting from
West Africa.
Almost nothing is known of the nesting behaviour of the
species. The sounds of nearby running water may help mask
any noise made by the birds during nesting.
TAXONOMY
The Bostrychia are very similar, and since few specimens are
available, much about their taxonomic relationships remains
to be understood. It was once suggested that the Hadada Ibis
and the Olive Ibis were the same species (Akeley in F.M.
Chapman 1912). As it is so much smaller, Bannerman (1930)
thought bocagei distinctive enough to be considered a different
species. The subspecies rothschildi is considered now to be
extinct and few specimens are available. The presently
recognised Bostrychia species were once separated into several
genera (Hagedashia, Lampribis and Bostrychia), and appear to be
related to the Madagascar Crested Ibis (Lophotibis) and the
Geronticusibises (Amadon 1953, Appert 1966, Fry etal. 1985).
Clearly, the taxonomy and relationships of the ibises in this
group need additional study.
CONSERVATION
This species inhabits deep forest areas that are rarely
visited, and as a result its status is almost completely un-
known. It is thought that one race is already extinct, and
another has only been seen once by biologists in recent
years.
Present dangers appear to include predation by forest
birds and mammals, especially baboons (Papio anubis, P.
cynocephalus], as well as the cutting of forest trees. On Sao
Tome and Principe Islands, deforestation is a major prob-
lem, especially since it appears that the birds are found only
in virgin forest. Dr P. Jones (pers. comm. 1990) reported
that the Sao Tome natives hunted the Olive Ibis in the last
century, when they were a 'favourite quarry'.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 315.
 Spotbreasted Ibis
Bostrychia rara (Rothschild, Hartert, and Kleinschmidt)

Lampribis rara Rothschild, Hartert, and Kleinschmidt, 1897. Novit. Zool., 4, p. 377: Denkera, Ghana

Other names:

Crested Ibis, Crested Wood Ibis (English); Kala (Congolese); L'Ibis Vermicule (French)

IDENTIFICATION
The Spotbreasted Ibis is a small (59 cm long) short-legged crested ibis confined to deep forests. It is a
dark-plumaged bird recognized by its crest and characteristically spotted neck and underparts.
The adult is brown with a long, loose crest, darker than the base colour and glossy green. The neck,
breast and underparts are reddish brown; the feathers are edged in dark brown with a central cream
spot. The mantle is also brown, edged in buff but with a glossy green sheen. The rump and tail are blue
green. The wings are a blue-black, the underwing black. The scapulars and inner secondaries sport an
iridescent bronze green. The bill varies between red and dark red, whereas the naked face and lores are
black, with a green to turquoise spot in front of the eye and another behind it. A green streak crosses the
face in line with the bill. The eye is dark brown; legs and feet are brown with a pinkish flush. Typical
measurements are: wing 27.0-29.0, tail 11.2-12.0, bill 11.5-13.0, tarsus 5.6-6.5 cm (Brown et al. 1982).
The species is sexually dimorphic, the bill of the female being smaller than that of the male. Brosset
and Erard (1976) noted that they observed the female to be less well marked on the head: the turquoise
spots and the facial streak were smaller, and the bill was not as bright red as the male. This singular
observation of sexual differences in soft parts may not be definitive as the female's courtship colours
may have started to fade before the male's. In any case, it would be expected that any difference in
markings would become inconsequential as nesting progressed.
Nestlings have red-pink skin, lightly purple-blue, and are covered with a sparse dark brown down.
The face is blackish blue and the chin pink. White down replaces the brown down after about 6 days,
when the sheaths of contour feathers begin to appear. The base of both mandibles are red. Immature
birds are similar to adults though with shorter bills, less pronounced markings and no metallic sheen.
This ibis is, like other Bostrychia, extremely noisy, especially at dawn and dusk when leaving or
returning to roost, and even at night (Brosset and Erard 1976). It is quiet during the day, while feeding
and at the nest. Brown et al. (1982) described its raucous call as 'K-hah, k-hah'. The second note is
accented, as was pointed out by Chapin (1932a) and Bannerman (1953) but missed by Brown et al.
(1982). When flying, the call may be given with alrpost every wing-beat (Bannerman 1953). The call
differs from those of the other Bostrychia ibises. Ash et al. (in press) were able to identify this species in a
new area of Uganda by its call, which lacks the honking, goose-like quality of the Olive Ibis. A tape of
the Spotbreasted Ibis by C. Chappuis (pers. comm. 1990) presents one bird with a soft voice and a
second one much harsher, with both birds accenting the second, or a third, syllable. This species
clatters its mandibles during nest relief and when threatened.
The Spotbreasted Ibis is distinguished among ibises by the speckling on its neck, breast and
undersides. The sympatric Olive Ibis is larger, lacks spots on the face and has purple on its crest. The
Hadada Ibis, which lacks a crest and is more likely to be found in the open, is also larger. All these
marks might be apparent close by and in good light, but the Spotbreasted Ibis's wooded habitat and
crepuscular flying habits make it difficult to distinguish in the field from either the Olive or Hadada ibis.
A. Brosset (pers. comm. 1990) considers them to be almost impossible to distinguish in the wild

except by voice, and J.S. Ash (pers. comm. 1990) certainly
confirms this with respect to distinguishing the Spot-
breasted from the Olive Ibis in Uganda.
DISTRIBUTION AND POPULATION
This species is found in West Africa from Liberia through
Ghana, and from Cameroon and Gabon through Zaire to
western Uganda. The presence of the species in Uganda was
recently discovered by Ash et al. (in press), where they
found it on the banks of the Kirimia River.
The Spotbreasted Ibis is seldom seen and is considered to
be rare, although widespread. It is most often reported from
northeastern Gabon, but much remains to be understood
about the details of its population status or distribution.
ECOLOGY
The Spotbreasted Ibis occurs in lowland Guinean and
Congo forests, where it is seldom seen, and so it is not well
understood (Bannerman 1953, de Naurois 1973). In
Uganda, Ash et al. (in press) found it along the river bank
associated with Afromomum swamps. Bannerman (1953)
stated that it is a bird of small forest streams and wooded
swamps and avoids the larger river banks and forest clear-
ings. It is primarily, although not entirely, a bird of virgin
primitive forest areas subject to periodic flooding.
This ibis feeds in pairs or alone by forest streams and in
wooded swamps and pools. It Probes in mud and other soft
substrate, consuming aquatic snails, worms, beetles and
grubs (Bannerman 1953, Bouet 1955a). The birds are pres-
Spotbreasted Ibis 205
ent in an area year-round, and use traditional roosts for
sleeping (Brosset and Erard 1976).
BREEDING
This species nests alone in the thick forests in which it is
found year-round. It apparently always nests near or over
water within the forest.
Nesting takes place when water is rising or is high and
appears to correspond with peak rainfall periods. In Gabon,
these are September-December and March-May (Brosset
1974). Brosset and Erard (1976) found that the two periods
of maximum nesting activity coincide with the two rainy
seasons, with some minimal activity also in the short dry
season of January to February but not in the longer dry
period from June to August. They nest in September and
November in Zaire, also the time of the big rains (Lippens
and Willie 1972)* However, nesting can probably occur at
any time of year under appropriate conditions.
Brosset and Erard (1976) suggested that initial pair-
bonding may take place in the traditional roosting sites. As
it is also likely that members of the pair remain together
year-round, courtship would be expected to be perfunctory.
On arrival at dusk, the birds circle around, changing posi-
tions frequently, perching on the main branches of trees. 5-8
birds perch side by side, bowing by lowering the head and
bill with crest feathers erect. Pairs touch bills and mutually
preen.
The nest is a small, crude platform of twigs very similar to
those reported for other Bostrychia, but smaller than that of
the Hadada Ibis and bulkier than that of the Olive Ibis
(F.M. Chapman 1912:24, Brosset 1974, I.S.C. Parker 1982).
It is some 30 cm across and between 5 and 15 cm thick; the
lining is of dead leaves, fibre, roots or fresh bits of epiphytes.
One nest was made entirely of liana shoots. The first nest to
be found was brought to P. Herroelen (1955:187). It had
been built at a height of about 3 m from the ground near a
stream in secondary forest. It was made of short, dry
branches and lined with fibre and roots. Brosset and Erard
(1976) observed a nest that was 80 cm above the level of the
water in the centre of the mouth of a stream that was about
4 m deep during the rainy season (0.8 m in the dry season).
This nest was on the main horizontal stem of one of the
lateral branches of a leaning shrub located at the edge of an
area of dense flooded vegetation. Another nest was in a
shrub, 4.5 m above water at the edge of the Ivindo River,
and a further one was 6 m above the water in a marsh on the
Libumba, a tributary of the Ivindo. Other empty nests were
found at heights from 1.5 to 3.0m in areas subject to
flooding but which were dry during Brosset's and Erard's
(1976) visit.
Although the nest is not hidden, it is effectively screened
by thick foliage, and the building, laying, incubating and
nest-relief are all carried out in a secretive, quiet manner.
The normal clutch size appears to be 2. The egg is pale
green, spotted with small irregular rusty markings, lightly
glossed and fairly large—55 X 35 mm. Both sexes incubate,
incubation taking about 20 days. It appears that the female
broods the eggs at night, the male replacing her in the
206 Spotbreasted Ibis
morning and early afternoon. During incubation, except for
occasional head or wing tosses to ward off insects, the bird is
immobile with its neck folded back and its bill placed under
the wing. It takes advantage of the need to turn the eggs for
stretching and shaking.
If threatened, the sitting bird will stretch out its neck and
clatter its bill strongly. On three occasions, monkeys (Cerco-
pithicus neglectus) passed near the nest being observed. Twice
the bird flattened itself against the nest, remaining immo-
bile. The third time, the bird, having just risen to turn the
egg, adopted an upright position with plumage tight, neck
extended and bill pointing upwards. It remained in this
position for up to 2 min without moving. The nest observed
was robbed of its young 9 days after hatching, but it is not
known whether the predator was animal or human.
A. Devez (in Brosset and Erard 1976) described sexual
differences in nest-relief after hatching, when the mate
comes to feed the young and replace the sitting bird. A
female arriving will raise her plumes, lower her bill and
make clattering noises, inserting her bill under the sitting
male to encourage him to rise. Bowing the body slowly
causes the male to rise and bill clatter with slightly ruffled
feathers. The young are then fed by the female. Should the
male not then leave the nest, the female extends her neck,
raises both crest and plumage and clatters loudly with her
bill until he departs. When a female is sitting and is relieved
by the male, he will pass his bill across her neck and they
will face in the same direction, bill clattering simul-
taneously. Sometimes this ceremony varies, with the male
passing his bill through the neck feathers of the female.
Feeding of nestlings is by regurgitation. Young chicks are
stimulated by the adult's tapping of the chick's chin and
clacking of its bill. The young bird then taps the adult's bill
starting at the tip and moving upward to the base then
inserting it into the open bill of the adult, whose crest is
often raised. As the chick grows, the initial stimulation of
the adult is no longer needed. Chicks Beg food by making
hissing noises described as shrill but only faintly audible.
Indeed the whole ceremony, including feeding and nest-
relief, is done quietly and with minimum noise and move-
ment. It is apparent that all activity at the nest is extremely
discreet, and creates the minimum of attention (Brosset and
Erard 1976).
TAXONOMY
Little is known about the biology, and hence the taxonomy,
of this species. Its relationships with other species, espe-
cially in the genus Bostrychia deserve additional study.
CONSERVATION
The status of Spotbreasted Ibis is really unknown. It
appears to be more abundant than does the Olive Ibis,
which however is overall more widely distributed.
Hunters are known to collect young from nests, and it is
likely that native hunters, as well as predators such as
monkeys, present the major threats to the species, but data
are insufficient to establish whether it is under severe threat
other than from man. Any encouragement to local hunters
to capture young for collections at museums or zoos should
be resisted to avoid the destruction of this rare species. The
forest of the Spotbreasted Ibis is rapidly shrinking, so they
will eventually become vulnerable through loss of their
habitat.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 316.
 Madagascar
Crested Ibis
Lophotibis cristata (Boddaert)

Subspecies:
Lophotibis cristata cristata (Boddaert)
Tantalus cristatus Boddaert, 1783, Table Planches Enlum., p. 51; based on 'Courly hupe, de
Madagascar' of Daubenton, 1765-81, Planches Enlum., pi 841: Madagascar
Lophotibis cristata urschi Lavauden
Lophotibis cristata urschi Lavauden, 1929, Alauda, 1, p. 233: Ankarafantsika and 100km SE of
Majunga, western Madagascar

Other names:
Crested Wood Ibis, White-winged Ibis (English); Akohoala, Akohovohitra, Akoholahialu, Lampir-
ara (Malagasy); Coq des Bois, Ibis Huppe de Madagascar (French); Madagaskar-Mahnenibis
(German)

IDENTIFICATION

The shy Madagascar Crested Ibis is a medium sized (50 cm long) rather handsome white-winged ibis,
with a long neck, long bill and long crest. It is the largest bird in the Madagascar forest.
The basic body colour of the adult is a heavy rufous brown. The crest on the nape and back of the
relatively long neck has a metallic sheen, the colour of which differs between the subspecies. The chin,
throat and neck are brown. The wing is mainly white.
The bare face is bright red, including around the eye. The rather long bill is horn colour. The iris is
dark, and the tarsus and foot are red. The sexes are alike. Breeding colour changes have not been well
described, but it is likely that the head, feet and crest colours brighten during courtship, as is the case in
most ibises.
The crest of the nominate subspecies consists of long feathering of dark metallic green and white. The
subspecies urschi has a deeper reddish brown body colour, and its throat, neck and crest are a metallic
green mixed with orange yellow and less mixed with white (Langrand 1990). Individuals having
intermediate characteristics are known.
Nestlings are initially all white turning brownish. The bill is grey, darker toward the tip. The bare
skin and feet are pale reddish yellow. Juveniles have a paler body colour than the adult, speckled brown
on the white wings, and white on the tail. The head is very pale brown. The grey bill is shorter and
paler, darkening from the base. The skin about the eye is grey, the eye itself is brown (Rand 1936). The
feathers on the back of the neck are long and blackish blue, but they lack the metallic sheen. The legs
are a paler red.
The voice is usually heard at dusk or when birds are flying to roost, but also when alarmed. It is
described as a monotonous, subdued, toneless 'Goo, goo, goo' by Langrand (1990) and as 'Grooh' by
Appert (1966). A call was described as a loud, far-carrying, slightly raucous 'Work, work, work' by G.S.
Keith et al. (1974). Rand (1936) stated that the bird can sometimes be heard calling during the day,

and described the sound as a loud, rather creaky *Ack, ack'.
Appert heard a bird at the nest give a quiet *Kr'.
The brown plumage makes this bird inconspicuous on the
ground, especially in its forest habitat. However, the white
wings flash strikingly when the ibis is in flight. Rand (1936)
saw 2 or 3 birds flying to a tree top with their wings flashing
conspicuously in the brilliant moonlight. When flushed, it
often flies to a perch in a nearby tree. Or it may walk quietly
further into the woods.
In flight, it cannot be confused with any other species in
its restricted range, as its wide, white wings contrast with its
dark body. It is the largest living terrestrial bird in Mada-
gascar, similar in size to the distinctive Helmeted Guinea-
fowl (Numida meleagris).
DISTRIBUTION AND POPULATION
The Crested Ibis is endemic to Madagascar, where pre-
viously it was distributed throughout the island, except in
the dry country of the far southwest. The subspecies cristata
is found in the east and north. The subspecies urschi occurs
in the west and south. It is present throughout the year, at
least in some areas (Appert 1966). It appears to be an
entirely sedentary bird.
No estimate is available of population size. Rand (1936)
found the species to be common, but G.S. Keith et ai. (1974)
reported it to be 'distinctly uncommon' in 1972. The west-
ern subspecies remains widespread and common (Appert
1966, Turner in Collar and Stuart 1985).
ECOLOGY
A forest ibis, this species is found in every type of original
forest on the islands and is not necessarily restricted to wet
habitats. In the east cristata is found in humid forest, in the
wet undergrowth of the dense rain forests by small forest
streams. It has also acclimatized to slightly degraded forest,
being found in thick and shady areas in vanilla and oil palm
Madagascar Crested Ibis 209
plantations. The subspecies urschi lives in dry deciduous
forest habitat, from sea level up to 2000 m above sea level.
This shy bird feeds alone and, more usually, in pairs. The
birds forage on the forest floor, by Walking through the
undergrowth. They stop to Probe amongst dead leaves and
forest litter, in the moss and around tree trunks. They also
Peck at the surface picking up insects and millipedes from
among the dry leaves (Gill in G.S. Keith et al. 1974). They
particularly feed along wooded streams at low water or in
debris covering the dry stream bed.
Generally, the diet of the Madagascar Crested Ibis con-
sists of invertebrates and small vertebrates (M.E. Nicoll and
O. Langrand unpub. report). Of 13 stomachs of cristata
examined, 10 contained various insects, including beetles,
one centipede, one spider and three earthworms. In one
stomach of urschi,, Rand (1936) found two 100 mm long
lizards, one 250 mm long snake and various insects.
The Madagascar Crested Ibis roosts on strong branches,
and flies to the top of large trees when disturbed. It prefers
shaded woodland areas.
BREEDING
This species is entirely dependent on tall trees for nesting in
the forest. The breeding period is from September to Janu-
ary according to Langrand (1990), and certainly includes
November since a female ready to lay was taken at Tabiky
on 2 November 1929. Appert (1966) observed nesting in
January and in November. A juvenile was collected at
Marotony on 2 January 1931 (Rand 1936).
Unfortunately little is known of the courtship behaviour.
Appert (1966) saw one bird of a pair display on the ground.
The bird ruffled its head, neck and chest feathers, and held
its wings out pointing to the ground. It took up one or more
dry leaves and held them for a relatively long time, and also
played with them and dropped them to the ground.
The distinctive crest is presumably used in courtship
displays, but it is kept lowered most of the time, except
when disturbed or alarmed.
Pairs nest solitarily. The nest is made of long sticks, 1-
2 cm in diameter. It is a large structure, up to 85 cm in
diameter, the cup being about 30 cm wide and 8 cm deep.
There is a lining of leaves and fine sticks.
The nest is constructed in the shade high up in the fork or
on several branches of a tree, as many as 7-15 m up in rain
forest trees (Appert 1966). Appert (1966) found a nest in a
tamarind tree (Tamarindus indicus) which was in leaf at a time
when other trees in the area were bare.
The eggs are a matt white; they measure, on average,
41 X 58mm. The usual clutch is 2-3, and both parents
share incubation (Langrand 1990). The incubating bird sits
tightly on the nest when approached from below.
Young in the nest gave a Teh' call when encountering a
person there; they also raise their wings and bristle their
feathers. They remain in the nest for 20-30 days.
TAXONOMY
The genus Lophotibis^ whilst superficially resembling Bostry-
chia (= Lampribis, Hagedashia), continues to be treated by us
as a separate genus pending additional study.
210 Madagascar Crested Ibis
The fact that distinctive differences exist in populations
on the same island is somewhat surprising. However, the
two forms occupy different habitats, and both apparently
are entirely sedentary. The forms are separated by the wide
central savanna, which stretches down the centre of Mada-
gascar.
CONSERVATION
The Madagascar Crested Ibis was the only bird endemic to
Madagascar to be afforded legal protection, as of 1974 (G.S.
Keith et al. 1974). However, it is still widely shot, hunted for
food or captured, for payment of a small fee (W.B. King
1981). It is also threatened by forest habitat destruction,
including forest fires, as are much of the native fauna and
flora of Madagascar. The protection of forest habitat is
essential to the continued existence of this and other forest
species on the island.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 316.
 Sacred Ibis
Threskiornis aethiopicus (Latham)

Subspecies:

Threskiornis aethiopicus aethiopicus (Latham)

Tantalus aethiopicus Latham 1790, Index Ornith., p. 706: 'Aethiopia' = PEgypt (Bruce, 1790, Travels
Source Nile, 5, p. 172, pi. 35)

Threskiornis aethiopicus bernieri (Bonaparte)

Ibis bernieri Bonaparte, 1855, Consp. Gen. Avium, 2, p. 151: Madagascar

Threskiornis aethiopicus abbotti (Ridgway)

Ibis abbotti Ridgway, 1893, Proc. US Natl. Mus., 16, p. 599: Aldabra Island

Other names:
White Ibis, Black-necked Ibis, Sticklebill (English); Heilige Ibis (Dutch); Heiliger Ibis (German);
Ibis Sacre (French); Ibis Sagrado (Spanish); Helig Ibis (Swiss); Ibis Sacro (Italian); CBHineHHbiH
Hbnc: "Ibis" (Russian)

IDENTIFICATION
The Sacred Ibis is a medium-sized, white ibis, 65-75 cm long, with black highlights, including its head
and neck, wing-tips and ornamental plumes along its back.
The adult's head and neck are bare and covered with a black scaly skin. The iris is brown and the
lower eyelid pale pink. The downcurved bill is black, and rather heavy looking for its size. Grey grooves
are quite apparent along either side of the mandibles. The distinctive plumes originate from the
scapulars and form a loose lace-like net across the back, closed wing and tail. When new, the black-
tipped primaries and secondaries are glossed with purple-blue and sometimes with iridescent blue-
green. A dull grey bare spot at the sides of the breast extends as a strip across the leading edge of the
underwing. The bare skin of the legs and feet also are black.
The island subspecies are smaller than birds on the African mainland. The subspecies bernieri has a
light blue eye and less black on the wing-tips. The subspecies abbotti has a bright blue eye, black wing-
stripe and reduced feather pigmentation on the wing tips.
The sexes are alike in appearance, but males are larger than females. Brown et al. (1982) reported
that the bill-length of males is 162-183 mm, whereas that of females is 135-157 mm. Urban (1974a)
reported that the differences were not sufficient to tell the sexes apart in the field except when both birds
of a pair were together or when a displaying group formed.
In courtship, the black scapular plumes develop a metallic bluish sheen. The bare skin underneath
the wings and the similar patch next to the breast become a vivid red. The black skin on the head and
neck becomes glossy, and the black legs flush red or dark red. A red ring becomes apparent in the iris.
Urban (1974a) reported that the plumage of birds nesting at Lake Shala became tinged with yellow or
brown, and underparts generally become stained as breeding proceeds.
There are some differences between subspecies in courtship plumage and colour. The ornamental
plumes are more developed in the subspecies aethiopicus but are less so in the subspecies abbotti and
bernieri^ in which the black plumes are also less intense during nesting. The African subspecies also has a
neck-sac, apparently absent in the other two races, which is especially obvious when half-open bills are
pulsated during hot weather such as when the bird is on the nest.

Nestlings have short, straight bills which are pale white or
pink, as are the feet. They are covered with a white down,
but the head, neck and back are black. At 21 days the dorsal
and lateral surfaces are covered with a brownish black down
(Urban 1974a). Wing-feathers are black as they develop,
and legs become greyish and distinctively swollen.
Juveniles are whitish with black speckling or streaking,
especially on the head and neck. Their wing-tips lack metal-
lic gloss, and the scapulars tend to be greenish brown. The
iris is brown with a narrow, outer, white ring. The feet and
legs are dull grey to blackish and these darken as the birds
get older. The bill takes 3 months to develop fully. Loss of
neck feathering and acquisition of full adult plumage is not
complete until the third year (FJ. Jackson 1938).
The Sacred Ibis is generally a silent bird but sometimes
utters a harsh groan or croak, especially when in flight. In
antagonistic encounters, both sexes call and squeal. Urban
described these calls as 'Whoot-whoot' and 'Pyuk-pyuk' and
also a wheezing 'Hnhhh-hnhh'. The female gives a
'Whaank* call in soliciting copulation. Adults call to young
with a 'Turrooh' (Cramp 1977).
Like all members of the Threskiornis genus, these birds
have strong, powerful wing-beats, which they alternate with
gliding, achieving 38 km/h (Urban 1974b). The head and
neck are fully stretched, and they fly in loose V-formations
or in line astern. They usually fly low over water but can
soar very high over land. R.A. Clark and A. Clark (1979)
report them travelling up to 30 km away from colonies when
feeding. The ontogeny of landing behaviour has been stud-
ied by G. Riippell (1977).
The Sacred Ibis is rather easily distinguished from other
ibises in its range by its white plumage, black head and
Sacred I bis 213
heavy bill. In flight it can be distinguished from herons and
egrets by its quicker wing-beat, and from storks because its
legs are hidden beneath its tail.
DISTRIBUTION AND POPULATION
The Sacred Ibis is a widespread resident in Africa south of
the Sahara desert, from Senegal in the west to Ethiopia in
the east, and southward to the Cape. Outside Africa, it
breeds only in Iraq, especially in southern areas of Amara to
Fao, although definite nesting records are few (Allouse 1953
in Cramp 1977). It was formerly found in Egypt, from
which country the type specimen and English name came,
but it had almost completely disappeared from there by
1850 (Meinertzhagen in Cramp 1977).
The subspecies bernieri is only found on Madagascar,
especially on the west of the island between Port-Berge and
Moromoe. The subspecies abbotti is endemic to the islands of
Aldabra, including West and Middle islands and lie
Polymnie.
African birds move around seasonally. There is a definite
movement of birds towards the equator in Africa, with
northern birds moving northwards and birds south of the
equator moving southwards at the commencement of breed-
ing, but movements vary depending upon seasonal rainfall.
Ringing records from South Africa show that birds ringed in
that country were recovered mostly in Zambia, with fewer
in Botswana and Namibia. The furthest travelled 1336km
(Dowsett 1969, R.A. Clark and A. Clark 1979). Birds in
Iraq are present year-round but probably also wander to
Kuwait and Iran.
After breeding birds disperse, moving in the dry season
especially to coastal areas and permanent bodies of water,
travelling considerable distances to traditional winter feed-
ing grounds. Numbers in these areas fluctuate seasonally
and annually. Neither of the island subspecies show signs of
either dispersal or migration.
The species occurs rarely in southern Arabia. Vagrants
have appeared in the USSR on the Caspian and Black sea
coasts, and 7 birds were seen in 1944 south of Baku, and it is
thought that they may breed in Khuzestan (Dementiev and
Gladkov 1951, Cramp 1977). It is possible, however, that
the birds seen in these areas were Oriental White Ibises.
Bururlin (1935 in Dementiev and Gladkov 1951) considered
them to be 'Indian ibises' (Oriental White Ibis), but
Dementiev disagreed, having seen them at close quarters.
That Eurasian Spoonbills migrate annually between the
Caspian Sea and Bharatpur in India raises the possibility
that Oriental White Ibises might do the same.
Few population estimates exist, but the species is com-
mon over much of the area in which it occurs. On the
Witwatersrand in South Africa, numbers fluctuate between
4900 in summer and 2500 in winter (Dowsett 1969, Tarbo-
ton 1977b). Elsewhere, feeding flocks vary from several
hundred birds to groups of 10 or 12, or even single birds.
They roost in the hundreds on islets in rivers or floodland,
on trees near water and sometimes even in villages. Their
dispersal in small parties make assessments of numbers
difficult over their large African range. Bernieri is uncommon
214 Sacred Ibis
(Langrand 1990). Abbotti numbered 150-200 in 1967-68
(W.B. King 1981).
ECOLOGY
This species is found in many habitats. It occurs in all types
of freshwater habitat and often feeds on dry land in grass-
lands and cultivated fields. Particularly in the dry season, it
frequents coastal mudflats. On Lake Turkana in northern
Kenya, we have seen birds feed on the rocky islands and
shores around the lake. Individuals wander some distance
into the desert areas where they feed on insects. At Lake
Naivasha, Kenya, they feed on hotel lawns, and along the
muddy shores of the lake. We have seen individual birds
feeding alongside Brown-necked Ravens (Corvus corax
edithae) scavenging at camps and villages. In South Africa,
they feed in sewage, dung heaps and refuse, as well as
cultivated fields, flooded grasslands and marshes (R.A.
Clark and A. Clark 1979). Abbotti feeds in coral pools and
along lagoon shores (W.B. King 1981).
They respond quickly to rainfall or flooding of potential
feeding habitat, moving into such areas immediately after
water rises (R.A. Clark and A. Clark 1979). Such flooded
feeding areas seem to be critical for the initiation and suc-
cessful completion of nesting.
The Sacred Ibis has simple methods of feeding. It will
Peck at the surface for insects on dry land, and Probe or
Deep Probe for crustaceans and aquatic insects in mud and
soft soil. Deep Probes can result in the bill being buried up
to the eye. They will Probe into carrion. Occasionally they
will run forward with wings half open to catch a flying insect
or moving reptile. They will also follow locust or grass-
hopper swarms.
Although they do sometimes feed alone, Sacred Ibises
more typically feed in groups. In shallow pools, they are
often found alongside African Spoonbills and Yellowbilled
Storks. They follow Little Egrets (Egretta garzetta) feeding on
prey that the egrets disturb (F.T. Morris 1978).
The diet consists of a wide variety of prey, especially
insects. It includes grasshoppers, locusts, crickets, water
beetles, worms, molluscs, crustaceans, fish, amphibians and
lizards. The birds consume the eggs of waterfowl, Great
White Pelicans (Pelecanus onocrotalus) and crocodiles (Crocody-
lus niloticus) (especially after the shells have been broken by
vultures), and the young of Cape Cormorants (Phalacrocorax
capensis) (Bolster 1931, Urban 1974a). Carrion and offal
figure in the diet near human habitation. R.A. Clark
(1979c) reported that South African birds in Pretoria con-
sume mainly human, animal and vegetable refuse, and
insects. In summer, the diet is 50% animal food and 34%
vegetable food; in winter, 47% vegetable food and 34%
animal food (by wet weight consumed). Urban (1974a)
found regurgitated food from a 1-month-old chick contained
22 dung beetles, 2 lepidopteran larvae, 2 coleopteran adults,
1 bead, 1 small piece of bone, 1 seed, bits from a millipede,
several small ants, and a small piece of egg shell.
Birds use communal roosts that are located in trees or on
islands. They arrive in the evening singly and in flocks
(S.M. Evans et al. 1981). These roost sites shift frequently.
In Witwatersrand, South Africa, only 3 of 28 roosts were
used year-round (Tarboton 1977a).
BREEDING
This species nests colonially in many different types of sites.
We have seen them nesting in trees and bushes, on the
ground and among rocks on islands. The species is surpris-
ingly resourceful in this respect.
Sacred Ibises will nest in single-species colonies or in
mixed colonies of storks, herons, spoonbills and cormorants,
but usually they choose to nest in discrete groups, somewhat
separate from other species. Colonies range from a few birds
to as many as 2000 pairs at Wembere, Tanzania (Britton
1980) and on the Witwatersrand in South Africa 2180 pairs
nested in six colonies in 1974 (Tarboton 1977a).
The breeding season varies geographically (E.K. Urban
1974a, pers. comm. 1988). Breeding is generally associated
with wet-season inundation, and often occurs at the height
of the local rainy season. Birds do even nest at the edge of
the desert in the wet season. They can nest in a single site
more than once in a year. We have seen up to 100 pairs
nesting in a mixed colony on the Tana River, Kenya, in
June during the long rains and 10 pairs in the same colony
during the short rains in January (Hancock 1984). In some
years nesting occurs at unusual times in response to unusual
flooding. In Madagascar, bernieri lays eggs in November
(Rand 1936).
Often nests are placed very close together, but they are
seldom touching. Nests are made of large sticks and lined
with softer material, mainly vegetation. They usually are
about 10cm across and 20cm deep, but sometimes much
larger, depending on the availability of nesting material.
When this is in short supply there is much stealing of sticks
and other material from nearby nests of other species in
the colony, as well as from each other. On Lake Turkana,
where sticks are scarce and the African Spoonbill nests be-
fore the Sacred Ibis, there is considerable fighting between
ibises and spoonbills as well as with Little Egrets
whose young have hatched before the Sacred Ibis begins to
nest.
The courtship behaviour of the Sacred Ibis has been
recorded by Urban (1974a) at a colony on Lake Shala in
Ethiopia. Nesting activity begins at the colony site as birds
fly in a Display Flight; these are male birds in small parties.
For the first few days they settle for short periods and
display by standing in groups with tail and ornamental
plumes spread and wings held down. The throat-sac of the
African subspecies is expanded. At this stage, the male birds
carry out a Sparring display, standing in a horizontal posi-
tion with head and neck extended and performing an open-
bill thrust at nearby opponents. As they strike the bills, they
make a wheezing squeaking call. The Sparring display is
performed by the displaying bird adopting a horizontal
stance with its head and neck stretched towards the op-
ponent, and with partially open wings revealing the bright
red skin. It then lunges with open bill.
Other displays at this time are the modified Sparring, in
which 2 or more males stand side by side facing the same

way and open their wings and point their heads skyward.
The Head Rub display is performed by the bird smoothly
moving its neck up and back and then, with bill pointed
upwards, flicking its neck backwards so that the crown
touches one side of the mantle. A Bill Popping display is
performed from a low horizontal position when the neck is
extended forward and down but the mandibles are not
snapped together. Males chase each other in pursuit flights,
and by supplanting attacks force opponents from their perch
by flying directly at them.
After 1 or 2 days the females arrive and constant chasing
and fighting occurs. If the female remains with a male and
no chasing occurs, pair-bonding usually commences. The
two birds face each other, in a Bowing display, with
downward-stretched necks and heads and intertwining of
necks and bills. The Head Rub and, infrequently, the Bill
Popping displays follow. Pairs then stand, with bodies
touching, on the nest territory which, when they have estab-
lished it, is used for copulation. The male stands over the
crouching female and grasps and shakes her bill, moving his
partially opened mandibles up and down it. After this, both
birds stand and preen, probably a Display Preen.
The male collects nesting material and the female re-
mains on site to build the nest. On the male's arrival with
material, the female crouches and both birds point their
bills upwards and intertwine them. When she has nearly
finished building, the female will utter a few sharp calls
(Brown etal. 1982).
Kim Lowe (pers. comm. 1990), in his study of South
African birds, considers that the behaviour of the Sacred
Ibis at the nest is considerably less aggressive and relatively
more silent than that of the Australian White Ibis. On the
other hand, Penny (1974) described a colony on Aldabra of
abbotti where fighting often occurs and the birds Tence with
their bills and kick each other like game cocks'.
Once the nest is completed, eggs are laid. Brown et al.
(1982) reported 2-5 eggs are laid, occasionally 5 in Ethiopia
and East Africa. The mean (of 535 clutches) in South Africa
was 2.43; in southwestern Cape 2.4 (291 clutches); in Trans-
vaal 2.0 (271 clutches). On Aldabra, Penny (1974) reported
2 eggs were laid, rarely 1 or 3.
The eggs are oval or slightly rounded, rough and dull
white with a faint blue tinge. A few dark red spots are
sometimes visible on African eggs, but Penny (1974) stated
that, on Aldabra, eggs were all-white. According to Urban
(1974a) incubation takes 28-29 days. A replacement clutch
after egg loss may be laid but this is not proven (Cramp
1977).
Both sexes incubate, changing over at least once every
24 h. Incubation commences after the clutch is complete
and hatching is synchronous (Urban 1974a, but see Stro-
nach 1968). The young are fed by regurgitation by both
parents, one of which remains at the nest for the first 7-
10 days. Young leave the nest between 14 and 21 days after
hatching and assemble in groups nearby. Adults continue to
feed them once a day until they fledge at 35-40 days. They
leave the colony after 35-48 days. Birds do not acquire full
adult plumage for up to 3 years. It is not known when they
first commence breeding.
Sacred Ibis 215
Nesting success is extremely variable. Manry (1978b)
reported 85% success from 54 nests on Schaapen Island,
South Africa. J. Parsons (1977) reported 43% from 353
nests at Kisumu, Kenya. Only 18 fledged from 420 chicks at
Kisumu in 1977. Factors affecting success include low rain-
fall affecting food supply, heavy rain storms causing young
to fall from the nest, and predation by African Fish Eagles
(Haliaeetus vodfer) (J. Parsons 1977) and Kelp Gulls (Lams
dominicanus) (Manry 1978b). Parsons reported Fish Eagles
eating 7 eggs per visit, and Manry reported the eating of
eggs, young and fully grown chicks by Kelp Gulls. On
Aldabra, disturbance by man caused desertions (C.W. Ben-
son and MJ. Penny 1971) and, in Africa, low-flying aircraft
have had the same effect.
TAXONOMY
We consider Threskiornis aethiopicus to be one of three species
forming a superspecies that includes T. melanocephalus and T.
moluccus. We thus follow Snow (1978) and K.W. Lowe and
G.C. Richards (1991) but not Holyoak (1970). These three
species are widely distributed in the Old World, and there
are difficulties in deciding the specific limits in this group.
Snow (1978) pointed out that two of the forms are them-
selves dimorphic, indicating that the clearest delimitation of
relationships would be by the recognition of three species.
K.W. Lowe and G.C. Richards (1991) found no evidence of
intergradation among the species in areas of overlap.
The Sacred Ibis has two isolated island populations,
bernieri on Madagascar and abbotti on Aldabra. K.W. Lowe
and G.C. Richards (1991) detected little difference between
these two forms. Holyoak (1970) stated that abbotti has little
or no black in the wing-tips and has duller display plumes
than bernieri. He also described the eye of the abbotti as light
blue, and bernieri as nearly white, though pale blue is the
more common description of the eye of the latter. In any
case, they both differ from the nominate race, which has a
brown to brownish red eye. Pending more details and in
view of the wide geographical separation of these sedentary
forms we prefer to treat them as two races. Interestingly,
aethiopicus has once been recorded from Aldabra, where it
was distinguished from the local subspecies abbotti by its
black wing tips and brownish-red eye (Penny 1974).
CONSERVATION
Despite its adaptability, this species is dependent on safe
nesting sites and on appropriate feeding areas. The African
subspecies migrates or disperses widely in search of the
latter and, except where pollution is present, finds ample
areas in which to feed. Nesting sites are, however, subject to
disturbance, drainage and predation by man.
The two islands subspecies are also under greater press-
ure, mainly from predation and disturbance by man. The
small population still resident on Madagascar also needs
careful protection as eggs and young are reported as being
collected by local people (Langrand 1990).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 317.
 Oriental
White Ibis
Threskiornis melanocephalus (Latham)

Tantalus melanocephalus Latham, 1907, Index Ornith., p. 709: India

Other names:

White Ibis, Oriental Sacred Ibis, Oriental Ibis, Black-headed Ibis, Indian Black-necked Ibis,
Oriental Black-necked Ibis (English); Witte Ibis, Zwartkop Ibis (Dutch); Didhar (Hindi); Hei-tou-lu
(Chinese); Dahakatti-Kokka, Thalkaththi-Chondam (Sri Lanka); Ibis a Tete Noire (French); Kuro
Toki (Japanese)

IDENTIFICATION
The Oriental White Ibis is a medium-sized (65-75 cm long), white ibis with a prominent black head
and neck, and a long downcurved bill.
The base colour of the adult is a pure white with bare orange patches under each wing and along the
flank. Like the closely related Sacred Ibis, the Oriental White Ibis has a featherless head and neck that
is covered by black skin. Its long bill is also black. Tom Roberts (pers. comm. 1990) reports a few
scattered blue spots on the head in winter. The eye is reddish brown. The legs and feet are dull grey to
black.
The sexes are alike in plumage. Sizes differences between the sexes are yet to be documented.
With the onset of the courtship season, a slate grey colour develops on the scapular plumes, and the
secondaries become elongated and frayed. These, and straw-like feathers that develop on the neck, are
the courtship plumes. The bare patches along the flank and under the wings turn a bright blood red.
The bare skin on the head and neck develops a bluish tinge. The feet and legs turn glossy black, flushing
red or dark red for a short period. The tail and axillaries are a pale yellowish brown.
Nestlings are thinly covered by white down, but the head, neck and back are black. The bill is black
and quite short. At 21 days the dorsal and lateral surfaces are covered in blackish brown feathers. The
legs at that stage are grey and become distinctively swollen. In the juvenile, only the grey face and
around the eye are bare. The rest of the head and neck are white-feathered with brownish or blackish
markings. The iris is brown and the bill shorter than in the adult. Juveniles lack ornamental plumes.
Full adult plumage takes up to 3 years to acquire.
The Oriental White Ibis usually is a silent bird. In the breeding season, however, it gives a
remarkably loud booming call (S. Ali and S.D. Ripley 1968). It also makes a series of grunts and
mumbles at the nest.
It flies with rapid wing-beats in strong, steady flight with occasional glides, often in loose V-formation
or in lines. In flight it is faster than egrets or storks.
This species is distinguished from other ibises in its range by its white plumage and black head and
neck. It differs from other members of its group in several ways (Holyoak 1970, K.W. Lowe and G.C.
Richards 1991). Compared to the Sacred Ibis of Africa, its bill is less robust, it lacks a neck-sac, it lacks
black on its wing-tips, its display plumes are light grey rather than black, it has straw-like neck plumes,
and it has a yellowish tail. It is distinguished from the Australian White Ibis by its more robust bill
218 Oriental White Ibis
and lack of black on wing-tips and display plumes. Its white
base-plumage distinguishes it from the Strawnecked Ibis. It
is distinguished from the Oriental Crested Ibis by its black
neck and head and dark legs and feet.
DISTRIBUTION AND POPULATION
The Oriental White Ibis was a wide-ranging species found
across continental Asia from India and Sri Lanka, north
and east to Nepal, Burma, Thailand, and China northeast
to Heilongjiang, south to Nam Phan (Cochinchina) in Viet-
nam and southern Cambodia, through the Malay Peninsula
in Malaysia to Java and probably Sumatra in Indonesia.
The species' distribution is somewhat patchy within its
overall range. Perhaps it is most widely distributed in India,
where it breeds throughout the subcontinent. Andrew
Robertson (pers. comm. 1990) found it in no fewer than 24
localities from Corbett National Park, across Rajasthan, at
Sultanpur Jheel near Delhi, Bharatpur, and in Gujarat in
the Gulf of Kutch, the Little Rann of Kutch, and southward
through Maharastra and Karnataka down to the tip of
India. It is common in the protected national parks and
lagoons of Sri Lanka. On the east coast of India it is found at
various jheels and tanks and along the coast at Point Cali-
mere. It is also found in Assam, India, and in the Sundar-
bans, but it is rarer elsewhere in Bangladesh. There are no
breeding records for Burma. In Vietnam, Vo Quy (in D.A.
Scott 1989) reported 7 large colonies in mangrove and
Melaleuca forest. While 1 colony has been abandoned, at
least 2 or 3 new ones have become established more re-
cently.
Isolated populations exist in China. We have seen them
nesting in small numbers in the Zhalong Nature Reserve in
the northeast of Heilongjiang. Although its overall range is a
broad one (Etchecopar and Hue 1978), we have found no
trace of the Oriental White Ibis in many of the regions we
have travelled over and consider it as probably no longer
extant over much of the illustrated range in China. T.-H.
Cheng (pers. comm. 1990) supports our impression.
The species is a resident of the Malay peninsula. Nesting
is not yet reported for Sumatra, although it may occur in the
Banyuasin Musi River delta (Silvius 1985). In East Java,
ibises nest at Ujung Pangkah near the Brantas (MJ. Silvius
and W.MJ. Vergheugt pers. comm. 1990). On Pulau Dua
and Pulau Rambat we found nesting birds in 1970 (Han-
cock 1984). Since that time, two colonies only remain on
Pulau Rambat, and the ibises do not nest on Pulau Dua
(Allport and Wilson 1986). Birds are still found at Muara
Bubos and Clanok and from Sutolilo (D.A. Scott 1989).
The Oriental White Ibis is somewhat migratory or dis-
persive over the most northern parts of its range. The
northernmost population remains in Heilongjiang until
October before migrating south (G. Archibald pers. comm.
1990). Birds that migrate from Manchuria traditionally
wintered in southern and eastern China and in Japan.
Today it is only a rare visitor to Japan, from Honshu
southward. It is found in Hong Kong in small numbers each
winter, and also in the Malay Peninsula. There seems to be
very limited evidence elsewhere that regular migration takes
place.
Most birds do leave their breeding grounds and disperse
to feed along the coasts and rivers, sometimes a long way
away. They wander to any suitable feeding areas, deserting
them when drought or disturbance affects foraging.
Occasionally birds wander to the Philippines (as well as to
Japan as noted above) (Itoh 1986, duPont 1971). Non-
breeding birds wander to the Nepalese terai, and a few still
visit the lower Sind valley in Pakistan. The species is
occasional in summer in Hong Kong. It is a vagrant in
Sulawesi and Kalimantan, Borneo, where birds come into
contact with the Australian White Ibis (C.M.N. White and
M.D.Bruce 1986).
Population estimates are few, but the numbers and range
have become much more restricted in recent years. There
were recently 25 nests in the northernmost colony site
in Heilongjiang, China (G. Archibald pers. comm. 1990).
Little is known about colonies elsewhere in China.
Probably the largest concentration of this species today
occurs in Vietnam at the southern tip of the Mekong River
valley. Vo Quo (in D.A. Scott 1989) reports 800-1000 birds
in the Bee Lieu coastal marshes and 500-600 breeding at
Cal Nuoc at the southern tip of the Mekong. There are 2 or
3 new colonies at Nam Can, and there are 7 large colonies in
the mangroves and Melaleuca forests in this area.
The largest population in India is undoubtedly at Bharat-
pur, where D.A. Scott (1989) reported 389 birds in summer
and 50 birds in winter at the Keoladeo Ghana Reserve in
1984. However at this reserve in 1983, the annual census
showed no fewer than 1435 birds (Hancock 1984). Up to
100 were counted at the Ranganthiloo Wildlife Sanctuary,
south of Mysore, where we saw them in 1988. Elsewhere
colonies of up to 50 or 60 birds are found in many areas of
India and Sri Lanka, though at Kaliveti Tank in Tamal
Nadu, India, up to 230 birds were counted, and at Kullar

Sandai and Vembakottai in the same state 160 were
reported (D.A. Scott 1989). This contrasts sharply with
Pakistan, where the largest recorded concentration was 100
birds in Sind in 1965, now reduced to only a few birds in the
1980s (Tom Roberts pers. comm. 1990).
The species is scarce in Burma, though a few winter near
Rangoon. In the southern central plains of Thailand, 90
birds roosted at Wat Tarn En and 70 birds were reported on
passage at Ban Lung Jorm (D.A. Scott 1989). It is rare in
Formosa and Japan. Similarly, it is now very rarely seen in
Malaysia (P. Round pers. comm. 1990).
In Indonesia only very small colonies have been reported.
Although breeding colonies have not been found, large
flocks occur in southern Sumatra, where the population may
be 2000 individuals (MJ. Silvius and WJ.M. Verheugt
pers. comm. 1990). Silvius et al. (1985) found 700 birds
feeding in Sumatra Selaan. The species occurred widely in
Java in the early part of the century but only recently were
breeding sites rediscovered, at Ujung Pangkah near Bran-
tas, East Java (MJ. Silvius and WJ.M. Verheugt pers.
comm. 1990). Also on Java, we reported 20-30 breeding
pairs on Pulau Dua in 1970 (Hancock 1984). Elsewhere
along the east coast, only small numbers were found.
ECOLOGY
This wide-ranging, adaptable species is both an aquatic and
terrestrial feeder, both inland and along coastlines. It will
feed quite often in dry areas and will even scavenge in open
drains as well as in city streets where it is not molested. It is
primarily a bird of the lowlands. In Sri Lanka, it is found in
coastal lagoons, in swamps and, in the dry zone, in irriga-
tion reservoirs.
Usually the Oriental White Ibis feeds in flocks or small
groups, but sometimes it feeds singly. It has simple foraging
methods, Walking Slowly, Probing or Pecking at the sur-
face. It will turn over vegetation and adopt tactile feeding
methods in shallow water, often in the company or other
species. The ibises use their long, curved bills to Deep Probe
on coastal mud flats. They shift territories as the tide rises.
Fish, frogs, molluscs, insects, worms and small reptiles
make up the bulk of the diet, but some vegetable matter is
also consumed. Stomachs of small downy nestlings con-
tained pulpy olive-brown and green vegetable matter and
elytra of tiny blackish beetles about 5 mm long (S. AH and
S.D. Ripley 1968).
BREEDING
Nesting occurs in well-protected areas, often in traditional
colonies mixed with herons, storks and cormorants, but
sometimes separately. Nests are typically built in medium-
sized trees such as Acacia, Prosopis, Barringtonia and others,
that are standing in or near water. Nests are also found on
the top of partially submerged shrubs such as Zizyphus and
Capparis horrida (S. Ali and S.D. Ripley 1968). Where they
were left unmolested, we have seen nests in tall trees on the
edge of human habitation.
Amongst other species, ibises nest in small conspecific
Oriental White Ibis 219
groups, though the ibises' nests are usually above those of
egrets and cormorants. On islands in the Gulf of Kutch,
they sometimes nest on or very close to the ground often
with Indian Darters (Anhinga rufa melanogaster]. In Indone-
sia, they nest alongside Milky Storks. Certainly here, as at
nearly all Indian Sites where we have seen Oriental White
Ibises nesting, they have nested in trees already occupied by
other species. The availability of nest sites in mixed colonies
is to some extent dictated by other species within the chosen
site. As an example, desertion of a traditional site by Asian
Openbill Storks at Bharatpur in 1989, due to artifically
raised water levels, left the trees clear for other species.
Nesting takes place after the rains and varies according to
the monsoons. The species nests in December to April in Sri
Lanka. Eggs have been found in April-August in West Java.
The nesting biology of this species has been studied by
Hoogerwerf (1935), but insufficient information is available
to determine the differences in the courtship and nesting
behaviour of any of the species of Threskiornis. One thing that
is fairly certain, however, is that the Oriental and Austra-
lian White Ibises do not have the throat-sac of the Sacred
Ibis.
Oriental White Ibises commence breeding activity with
Display Flights, landing for short periods while they display
by standing in groups with tail spread, wings held down and
ornamental plumes spread. The blood-red stripe under the
wing is prominent in Display Flights.
Males carry out Sparring displays standing in a horizon-
tal position with head and neck extended and feinting with
an open bill thrust at opponents. No bill snapping occurs.
Circular pursuit flights are common at this stage. We have
seen, in this species as in the Sacred Ibis, a modified Spar-
ring display in which two birds stand side by side facing in
the same direction and point their bills upwards and, nearly
touching each other, call with a harsh grunt.
Male birds advertise for females by performing Bill Pop-
ping and Head Rubbing displays, and these are carried out
also by females as they approach the proposed nest sites.
Bill Popping- is the less common of the two. The bird keeps
low, crouching to extend its neck forward and down in a
sudden quick movement. In the Head Rub, the neck is
raised up and back with the bill pointing skyward and then
the head touches the mantle on one side. When a female
approaches a displaying male, either she is chased away or
performs a Bowing display. Both birds Bow with neck and
head extended, necks may be inter-twined, bills are crossed.
Birds will then Display Preen, but not allopreen. We have
found the Oriental White Ibis to be less aggressive than the
Sacred Ibis and to have less elaborate displays.
In fact, we have seen considerable variability in the inten-
sity and duration of courtship displays. There is less fighting
and less intense displays in small colonies, where only a few
pairs are present, than in larger ones. Our observations of
courtship in a year following highly successful breeding
suggested that breeding activities were particularly strong
then, whereas in a year after a nesting failure, displays were
not so extended (pers. obs. J.A.H.).
Once nesting pairs are established, the remainder of the
competing group will depart and re-form at another site,
220 Oriental White Ibis
although they may return to attempt to dislodge the first
group of paired birds.
Copulation takes place at the nest-site after the territory
has been successfully established. The male straddles the
crouching female and grasps her bill, shakes it, and then
moves his partially opened mandibles up and down her bill.
Preening follows. Copulation is carried out usually before
any nest-building is started. K. Lowe (pers. comm. 1990)
noted frequent promiscuity between unpaired birds as well
as between males already paired, which copulate with other
females visiting the nest-site.
Stick gathering is carried out by the male and the nest
built mainly by the female. S. Ali and S.D. Ripley (1968)
stated that both birds actually take part in nest building.
However, competition for sticks is usually such that one bird
remains at the nest, although it does steal sticks from neigh-
bouring nests whilst remaining on its own platform.
Nests are often close together, in some cases with only a
few centimetres separating them. Nests built close together
inevitably lead to aggressive action around them. Once the
nest is completed, one bird stays there.
Nests consist of a small platform of sticks c. 25-30 cm
across. 2-4 eggs are laid. They are long, oval-shaped and
chalky white with a faint bluish tinge, and sparingly marked
with small spots and blotches of light or dark brown, more
densely at the broad end. The average size of 150 eggs
measured was 63.5 X 43.1 mm (E.C.S. Baker 1929). Incu-
bation takes about 23-25 days.
Young are fed by regurgitation and both parents feed the
young. The chick will grasp the adult's bill to open it and
then thrust its bill into the adult's opened gullet. During the
heat of the day chicks are not usually fed, and often both
parents will stand on the nest to shield the chicks from the
hot sun. At that time the adult's bill is half open and the
throat of the bird pulsates continually.
There is constant raiding of the nest by House and Jungle
Crows. Young ibises are attacked by a variety of birds of
prey in India. Fledglings leave the nest to join other young
from surrounding nests and sit on branches close to one
another. At this time they will still food Beg when adults
pass over head. Seldom do more than 2 chicks survive per
nest.
TAXONOMY
The Oriental White Ibis has variously been considered a
separate species or a race of the Sacred Ibis (Holyoak 1970,
J. Steinbacher 1979, Van Tets 1978). However, K.W. Lowe
and G.C. Richards (1991) showed that morphological vari-
ations in museum specimens were sufficient to separate T.
melanocephalus from both T. aethiopicus and T. moluccus. They
found no intergradation of characters between moluccus and
melanocephalus, despite the close contact of the forms in the
Sulawesi/Moluccan island area. C.M.N. White and M.D.
Bruce (1986) consider their distribution to be allopatric.
K.W. Lowe and G.C. Richards were not able to document
detailed courtship displays, although suggesting that the
behaviour of the forms is strikingly different.
We consider that these forms have been isolated for a long
period and that it is best to treat them as separate species
(K.W. Lowe and G.C. Richard 1991, D. Holyoak pers.
comm. 1990). Certainly there is room for considerably more
study before the taxonomy is fully resolved.
CONSERVATION
Threats to this species include loss of habitat due to drain-
age, disturbance by human predation and pesticides. The
establishment of protected breeding areas in reserves, par-
ticularly in India and Vietnam, has helped the species to
maintain its numbers in these areas, but in most other
regions it must be regarded as a threatened species.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 318.
 Australian
White Ibis
Threskiornis molucca (Cuvier)

Ibis molucca Guvier, 1829, Regne Animal, nouv. ed., 1, p. 520: Moluccas

Subspecies:
Threskiornis molucca molucca (Guvier)
Threskiornis molucca pygmaeus Mayr
Threskiornis aethiopicus pygmaeus Mayr, 1931, Amer. Mus. Novit., no. 486, p. 6: Rennell Island,
Soloman Is.

Other names:

Australian Ibis, White Ibis, Black-necked Ibis, Sticklebill (English)

IDENTIFICATION
The Australian White Ibis is a medium sized, white ibis with a black bill, bare black neck and obvious
black plumes covering the lower back. This Australian bird looks very much like its relative, the Sacred
Ibis of Africa.
The adult's white base colour is set off by the dark, green-black wing-tips and black, lace-like
scapular plumes that extend over the lower back and over the yellow tail. The bare face is black to blue-
black; the bill is black. The iris is brown. A pink band traverses the back of the head. The neck is
covered by bare, black skin and lacks a neck-sac. The upper part of legs are pink, lower portions dark
grey to black. Patches of bare reddish skin occur at the sides and under the wings. Elongated feathers
form a tuft at the base of the foreneck.
The sexes have similar plumages, but males are larger than females. Some comparative measure-
ments for molucca are, for the male, culmen 16.8-19.0, wing 36.8-38.1, tarsus 9.1-10.0, tail 12.1-12.5 cm;
and for the female, culmen 13.9-15.1, wing 34.8-35.5, tarsus 8.3-8.5, tail 11.5-11.8 cm. The size of the
molucca varies along a cline, increasing from New Guinea through southern Australia (K.W. Lowe and
G.C. Richards 1991).
The subspecies are distinguished primarily by size, the island subspecies (pygmaeus) being considered
a dwarf form. In comparison to the data presented above on the Australian form, the body-part sizes of
a pygmaeus specimen are culmen 12.3, wing 30.4, tarsus 7.4 and tail 11.0 cm.
During courtship, the bare patch under the wing and flank turn bright red, and the other bare parts
become glossy black. Ornamental plumes reach their yearly maximum extent, and the tuft of feathers at
the base of the foreneck reaches maximum elongation.
Nestlings are at first covered in white down except for the head and neck, which have black down.
The white feathers are replaced after 3 weeks with brownish black feathers. At this time the legs are grey
and swollen. The juvenile is whitish but has black feathers on the head and lacks neck and wing
ornamental plumes. The bill is shorter and straighter than is the adult's.
Like the other Threskiornis species, this is generally a silent bird, except during courtship and at the

nest. The voice, as described by Pizzey (1980), consists of
harsh barks and shouts. The normal call is an 'Urk',
whereas calls associated with breeding are 'Koaha*, 'Taw-
taw' and 'Taw-aw' (Frith 1976). Young birds utter a
squeaking, hissing sound.
The Australian White Ibis is often found in company with
the similar Strawnecked Ibis but is easily distinguished by
its white body. Its small neck-tufts contrast with the full,
straw-coloured neck plumes of the Strawnecked Ibis. It is
distinguished from the remaining Threskiornis species by a
combination of characters including its dark wing-tips, dark
ornamental plumes, a neck-tuft, no neck-sac, yellow tail and
coloured tubercules on the head. In comparison with the
Sacred Ibis of Africa, it has a more delicate bill, lacks a
neck-sac and has a neck-tuft. Unlike the Oriental White
Ibis, it has black wing tips and display plumes.
DISTRIBUTION AND POPULATION
This species is widely distributed on the Australian main-
land north to southern New Guinea, the Moluccas and the
Solomon Islands.
In Australia, this ibis is particularly found in the east and
north. In the southeast it is a common breeder along the
coast and inland but not further than 3CFS. There are some
colonies in the northeast, and we have seen colonies on the
South Alligator river east of Darwin. Only since 1952 have
birds moved into the southwest region, and breeding there
was first reported in 1979. In New Guinea, the bird is
resident in Irian Java and Papua New Guinea. It also
inhabits islands of the Moluccas (Seram, Geser, and great
Kai). The subspecies pygmaeus is found on Rennell and
Bellona in the Solomon Islands.
Both migratory and nomadic, this species responds to
seasonal and localized dry periods by undertaking popu-
lation shifts (Carrick 1962). Bekle (1982) found it to be
somewhat unpredictable in its wanderings in the Perth area.
Banded birds from Victoria and New South Wales moved
northward along the east coast and to the Cape York Penin-
sula. Birds cross the Torres Strait to winter in Papua New
Guinea (Purchase 1976). Draffan et al. (1983) reported
migratory flocks of 20-500 birds crossing into southern New
Australian White Ibis 223
Guinea. Thus the resident population in New Guinea is
joined by migrants from Australia in winter.
The species is accidental in New Zealand and Tasmania
(W.R.B. Oliver 1955, R.H. Green 1959a). In Tasmania
there were major eruptions in 1957-58 and again in 1975,
but breeding has not been recorded. It is also vagrant in the
Lesser Sundas (Tinimbar and Babar), Moluccas (Kai,
Geser), Sulawesi, and Kilimantan on the Borneo coast,
where it may or may not coincide with the Oriental White
Ibis (K.W. Lowe and G.C. Richards 1991, MJ. Silvius and
W.J.M. Verheugt pers. comm. 1990).
Population sizes are in places fairly well known, at least in
Australia. The major colonies are found in Victoria and
New South Wales. In 1979-80, numbers in 16 colonies
ranged from 150 to as many as 15000 per colony, with a
total of 33000 birds (Cowling and Lowe 1981), and on
Barren Box Swamp in 1967 there were up to 1000 nests.
However, colonies in the north are smaller. There were an
estimated 184 nests at Kapalga and up to 1000 nests on the
Fitzroy and the Ord Rivers. We visited a colony site on the
South Alligator, which was also occupied by a mixed breed-
ing colony of egrets and cormorants. There we saw up to 50
nests of ibises, whilst, across the river from the main colony,
2 more colonies of up to 10 nests were found with no other
species present.
The Australian White Ibis is not uncommon at Seram
and has been considered to be scarce or uncommon in
western Irian Jaya. However, a recent survey in New
Guinea found large mixed colonies of nesting birds in south-
eastern Irian Jaya, in the Pulau Kimaam and Wasur re-
serves (Bishop 1984). Over 15000 Australian White Ibis
were recorded at Pulau Kimaam and over 30 000 at Wasur.
This exciting discovery alters the picture completely as to
the status of this species, as well as the status of other storks
and ibises in New Guinea.
ECOLOGY
The Australian White Ibis much prefers aquatic or wet
situations and tends to avoid arid areas (there are few
records from the centre and central south of Australia ex-
cept during exceptional rains). It feeds along the shallow
edges of lakes and inland lagoons, on mudflats, in swamps,
irrigated pasture land, mangroves, creeks, marshes, and at
dams, reservoirs and rivers. In times of drought, birds feed
along the coast. In Irian Jaya, birds occupy the rich grass-
lands that lie within the Wasur reserve and extend eastward
to join the extensive Bulla Plains of the Tonda Wildlife
Management area (Bensbach, Papua New Guinea) (Bishop
1984). In northern Australia we watched birds feeding
along the mudbanks of the heavily tidal rivers on the vast
expanse of exposed mud, and on the grassy flood plains.
Human habitats are not avoided. Birds commonly visit
garden lawns, playing fields and sewage dumps. They sca-
venge around garbage tips and poultry yards. Like others of
the genus they will also feed on burnt savanna.
Australian White Ibises feed in conspecific flocks or in the
company of other wading birds. We have seen them with
Royal Spoonbills, Pied Herons (Egretta picata) and Black-
224 Australian White Ibis
necked Storks. Whitefaced Herons (Ardea novaehollandiae)
follow Australian White Ibises, capturing prey disturbed by
their probing (W.E. Davis pers. comm. 1987).
The Australian White Ibis feeds by Probing, although it
also has been reported to Head Swing. A bird will wade
steadily through the water Probing near its feet (Vestjens
1973a). In doing so, it will often penetrate 25cm or more
into the water, deeply submerging its head and neck. This
behaviour contrasts with that of the Strawnecked Ibis,
which prefers shallower water (Carrick 1959). Australian
White Ibises also use 'anvils' for opening mussels. On
finding a mussel, the bird takes it out of the water, holds it
with the feet on thick roots of living trees, pebbles or a flat
rock and hammers it with the bill until the shell breaks
(Vestjens 1973a).
The diet is varied, consisting of earthworms, freshwater
crayfish, spiders (adults and egg-sacs) crickets, mole crick-
ets, grasshoppers, shieldbugs, moths and caterpillars, bees,
ground beetles, water beetles (adults), dung beetles, click
beetles (larvae), weevils, freshwater mussels, water snails,
fish, frogs, and lizards (Carrick 1959).
In eastern Australia, Carrick found that the White Ibis
fed largely on freshwater crayfish and other aquatic foods,
and made limited use of drier habitats. He also found that
this ibis favoured larger prey than the Strawnecked Ibis,
with most insects being consistently about 5 cm long; a large
proportion of the diet involved Probing for burrowing types
of prey such as crayfish, crabs and earthworms.
Cowling (1974) found that captive birds preferred to eat
live locusts rather than dead ones, but preferred sluggish
ones available in areas sprayed with insecticides. Among
grasshoppers and locusts (Acrididae), Carrick (1959) found
that the birds preferred the spur-throated locust (Austracris
guttulosa), which is abundant around the margins of marshes
after feeding, and that in many cases dry-land species were
consumed by Strawnecked Ibises but not by the Australian
White Ibis.
BREEDING
K.W. Lowe (1984) stated that, in Australia the White Ibis
exhibits two broad responses to its environment. Some
populations accommodate their breeding to an unpredic-
table environment and other populations use complex
breeding tactics dependent on a highly predictable set of
environmental conditions. Overall, the species breeds dur-
ing the wet season in locations where recent rains have
produced the flooding that sustains breeding.
Nesting is in trees, bushes or reeds. In Victoria, birds nest
in lignum (Muehlenbeckia cunninghamii), reeds (Phragmites
spp), rushes (Juncaceae) or eucalyptus trees (Eucalyptus). In
the north and in the south of the State, nest sites are usually
in reeds, rushes or paperbarks (Melaleuca ericifolia). Rarely
do Australian White Ibises nest on the ground, though they
do so occasionally on islands on the perimeter of wetlands.
They have been recorded nesting in trees on dry land well
away from wetlands (Cowling and Lowe 1981).
This species often nests in close proximity to the Straw-
necked Ibis in Australia. In New Guinea, it nests alongside
herons, egrets, cormorants and storks. Here we have seen
that it has a tendency to nest in groups of its own species,
and sometimes in separate colonies at some distance from
the mixed colonies.
Courtship behaviour has not been well documented, but
some information is provided by Frith (1976) and K.W.
Lowe (1984). Males and females gather in 'courting
groups'. Courting groups disperse and reform elsewhere.
The nest is started when groups have dispersed, leaving a
pair that has mated to establish it. Within a group, synch-
rony of breeding is established, but sometimes other groups
will take over occupied nest-sites, evicting the pairs and
destroying the nests, which may contain eggs or young.
Males display by vocalizing, erecting feathers, spreading
their wings and snapping their bills. Raising their wings
displays the scarlet skin underneath and alongside the
breast. The male chooses a potential nest-site and tramples
a small area. The male uses Bowing displays, and when a
female approaches, the male grasps a twig in its bill. Both
birds will grab the stick together, after which they Display
Preen.
Fighting is common. When one male was seen to inter-
rupt the display of another, they fought by Sparring with
their bills. Fights occurred between paired and unpaired
males over the possession of a nest-site, or when an unpaired
male or female attempted to intervene between a female and
her male 'escort'. Females that fight for access to a male are
usually individuals that have bred with the same male
previously. These fights occur during courtship bouts and
on nest-sites after a pair had become established. Fights
between females have resulted in eggs being destroyed by
the interloper.
The male and female share the task of nest building. A
nest consists of a substantial, compact, shallow cup of reeds
or sticks lined with vegetation. In dense colonies, nests
become trampled into a continuous platform (Pizzey 1980).
The Greeting displays at the nest consist of deep bowing.
Promiscuity is common both before and after pair-bonding,
and some birds make several attempts to breed during the
season.
Between 1 and 4 eggs are laid, most pairs having 3 eggs,
which are dull white and sometimes become heavily stained.
Eggs are laid at night on alternate days. The mean size of
eggs is 67.8 X 43.8mm (n = 30) (K.W. Lowe 1984). Both
sexes incubate, incubation being 20-23 days.
Hatching is asynchronous, over a 2-day period. Feeding
of nestings is by regurgitation. Young remain in the nest for
approximately 3 weeks, and then stay together in groups for
another 2 weeks, perching together on trees while still being
fed by the parents.
The mortality rate of chicks depends largely on avail-
ability of food. However, colonies are often subject to flood-
ing and are deserted. Conversely, in areas where sites dry
out, nests are also deserted through lack of food availability
or because of land predators. Predation does not seem to be
as important a factor as it is for the Sacred Ibis in Africa.
We have, however, seen birds of prey circling nests in the
South Alligator colony site, and eggs and young are cer-
tainly taken by Whistling Kites (Haliastur sphenurus), which

also harry adult birds taking food to the nest (Pizzey 1980).
Lowe (1984) found the number of young fledged to be
between 0.50 and 0.60/clutch.
TAXONOMY
The forms of the Australian White Ibis have in recent years
usually been considered to be representatives of the Sacred
Ibis (J. Steinbacher 1979). However, more recent consider-
ations have concluded that the Sacred group is best classi-
fied as three species, which we follow (Snow 1978, Mees
1982, K.W. Lowe and G.G. Richards 1991).
The aethiopicus and molucca species resemble each other
superficially in plumage colour. However, it is likely that
molucca and melanocephalus are more closely related, sharing
such features as the tuft of feathers on the breast and
juvenile plumage (Snow 1978). K.W. Lowe and G.C.
Richards (1991) show that there is no blurring of character-
istics between molucca and melanocephalus, despite the close-
ness of their ranges. Three subspecies have been recognized
in recent years within the molucca group, molucca, pygmaeus
and strictipennis. The subspecies molucca represented the birds
in the Moluccas, and strictipennis the birds in Australia. The
birds in New Guinea were considered to be intermediate (J.
Steinbacher 1979). Holyoak (1970) had distinguished be-
tween the two based on the colour of the shafts of the
secondary wing-feathers, but Lowe and Richards show this
character to be variable among birds and probably among
Australian White Ibis 225
ages. Mees (1982) and K.W. Lowe and G.C. Richards
(1991) concluded that molucca and strictipennis are identical.
However there is a geographical variation in size, decreas-
ing from north to south.
The name molucca has priority for the species name. The
name moluccus is often used, but Sibley and Monroe (1990)
show that the original name, molucca, is correct.
The subspecies pygmaeus is similar to molucca but is a dwarf
form, being considerably smaller than the nominate race.
Thus we consider that the Australian White Ibis is a polyty-
pic species.
CONSERVATION
This species flourishes in suitable habitat throughout its
range and is to our knowledge, in no difficulty. Recent
discoveries in New Guinea are especially encouraging. Re-
serves in Australia, at least, are well protected, and the
species is not subjected yet to the pressures of human popu-
lations, either there or in Irian Jaya. Furthermore, it
appears to accommodate well to human habitations.
Note: Body measurements, egg measurements and nestingne
season data for this species can be found in the Appendlix on
page 318.
 Strawnecked Ibis
Threskiornis spinicollis (Jameson)

Ibis spinicollis Jameson, 1835, Edinburgh New Philos. Journ. 19, p. 213: Murray River, New South
Wales

Other names:

Dryweather Bird, Farmer's Friend, Letterbird, Black Ibis, Black-and-white Ibis (English)

IDENTIFICATION
A medium-sized ibis, 65-70 cm long, the Strawnecked Ibis is a predominantly black and white bird,
with straw-like display feathers at the base of its neck.
The adult is in many ways similar to the more widespread Australian White Ibis, with which it is
often found. The back and wings are black with a glossy green sheen. Part of the neck, the undersides
and tail are white, which provides a conspicuous contrast when the bird is in flight. Just below the head,
at its side and base, the neck turns predominantly white, merging with the stiff straw-coloured plumes
protruding from the lower neck. A stripe of white feathers crosses the undersides of the wings.
The unfeathered head, back of the neck, and throat are black, as is the long, curved bill. The feet and
lower legs are black, but the upper legs are paler, flushed pink to red. A patch of bare black skin occurs
on the flanks.
Males are larger than females. McKilligan (1975) found the length of the culmen (straight line from
tip to base) averaged 165mm in males contrasted with 137mm in females. This difference was
statistically significant and could be used to determine the sex of birds at close range in the field.
During courtship and breeding, the legs turn pink to dark red, the green sheen brightens and the
straw-coloured plumes are at their maximum length.
Immature birds have feathered heads, and overall are duller coloured than adults and lack the sheen
on their black feathering. The chin and the back of the neck are black, the legs and feet dusky. The head
feathering is lost and neck plumes develop in the third or fourth year (Le Souef 1917).
The Strawnecked Ibis is generally a silent bird. Its voice has been described as consisting of grunts,
hoarse coughs, and drawn-out croaks (Pizzey 1980, Simpson and Dat 1984).
In flight this bird is a study of black and white contrast. Flocks will wheel and turn in loose formation
often descending from great heights, circling widely toward the ground in a series of gliding movements.
The Strawnecked Ibis is distinguished from its two sympatric ibises, the all-dark Glossy Ibis and the
Australian White Ibis, by its colour pattern. It is less confined to wetlands than are these species.

DISTRIBUTION AND POPULATION

The Strawnecked Ibis has long been considered to be an Australian endemic. It nests at least
occasionally in all states and territories of the mainland, although it is usually absent from the arid
interior (McKilligan 1975). The population appears to be centred in Victoria, with fewer nesting
elsewhere in the continent. The nesting populations in western Australia and in southeastern Australia
are separated by deserts.
Recently the Strawnecked Ibis has been found breeding in southeast Irian Jaya, which considerably
extends the breeding range of the species (Bishop 1984). Prior to this discovery, the species was
228 Strawnecked Ibis
known to be found in New Guinea nearly year-round, but in
varying numbers. It was reported over 100 years ago by
D'AIberis at Attack Island, Fly Delta, where in December
large flocks were seen flying at a great elevation in a north-
west direction. It was found throughout the year in the
Bensbach River area of the southern trans-Fly region. Its
abundance there varied with rainfall, peak numbers being
reached at the beginning of the rainy season, when several
thousand were present, the numbers falling to hundreds at
the height of the dry season. It was also seen fairly regularly
by Hoogerwerf at Kurik in the nearby south of Irian Java,
but seldom in the rainy season. Clearly, much of the move-
ment of birds into and around New Guinea is migratory. It
has in fact long been known to be a migrant to Irian Jaya,
and flocks of 50-100 birds are seen in the dry season over the
Torres Strait (MJ. Silvius and W.J.M. Verheugt pers.
comm. 1989). Flocks have been found drowned at sea (Pes-
cott 1968).
Such a migratory dispersal appears to be the rule, espe-
cially northward from the densely packed breeding areas in
southeastern and western Australia.^ Birds nesting in New
South Wales migrate to the east coast and the northeastern
Northern Territories, whereas birds nesting along the west
coast migrate north to Darwin and Arnhem land. We have
seen flocks arriving in April, apparently from great dis-
tances and heralding the onset of the hot, dry season there.
The arrival of the Dryweather Bird, landing among the
Magpie Geese, Anseranus semipalmata, with their young is the
harbinger of the season to local residents.
Immature birds disperse widely and rapidly after the
nesting season. These birds wander widely, especially in
response to changing water conditions, occurring at times
nearly anywhere in Australia. The species is a vagrant in
Tasmania and in Indonesia, beyond New Guinea.
Of the three species of ibis found in Australia, this is
the most numerous. In 1979-80, 76500 adult Straw-
necked Ibises were estimated to have bred in the 26 largest
colonies in Victoria (Cowling and Lowe 1981). Other docu-
mented nesting concentrations are 150000 birds in most
years at Bool Lagoon in South Australia, 200 000 at Reedy
Lake, near Kerang, Victoria, 100000 on the Lachlan River,
near Booligal, New South Wales, and 15000 on the Yanko
Creek, New South Wales (McKilligan 1975, Cowling and
Lowe 1981). In earlier years, even larger colonies have
been reported (Waterman et al. 1971). The largest colonies
in the southwest are a few hundred birds. The newly dis-
covered nesting population in Irian Jaya included about 350
birds.
The Strawnecked Ibis remains abundant throughout its
range in Australia, with no indication of population de-
clines. In Victoria, Cowling and Lowe (1981) did not detect
a change in numbers of nesting ibises over 25 years. How-
ever the breeding range appears to have expanded towards
the south-central and southeast of that State. The species
was rare in southwestern Australia, until 1982.
ECOLOGY
The Strawnecked Ibis is more likely to be found in dry areas
than the other ibises in Australia, often moving into dry-
land territory. It seems particularly attracted to insect pla-
gues in dry areas, although there is no evidence that it can
control outbreaks. It also follows grassland fires (M.J. Sil-
vius and W.J.M. Verheugt pers. comm. 1989). The unpre-
dictable climate of Australia means that the Strawnecked
Ibis is often compelled to assemble in wandering nomadic
flocks to use these sporadic food sources.
Among its typical foraging habitats are grasslands, in-
cluding city parks and road edges, swamps, and cultivated
land after the crop has been harvested, but birds have not
been seen in long, dense grass or in woodland with thick
undergrowth (McKilligan 1975). They also avoid marine
habitats. Use of dry habitats becomes especially prevalent
during the non-breeding dry season, in the North. During
this period the ibis chooses sites of moderate to average
moisture content rather than deep water or croplands.
Unlike the Australian White Ibis, the Strawnecked Ibis
does not usually visit garbage dumps.
Feeding in wetland areas is by deep or shallow Probing
along the edges of marshes and in wet grasslands. Here the
birds are more likely to keep some distance from one
another, but when they move to terrestrial feeding areas
they move in closer formations. When grasshopper and
other swarms of big insects are found, the ibises feed by
sight in large close parties moving rapidly forward. They
feed more by visual techniques than do Australian White
Ibises. They also have been reported to feed in trees, by
probing into cracks in the bark (Haffenden 1981).
The food of this species has been well studied by Carrick
(1959), Cowling (1974), and McKilligan (1975). Their diet
predominantly includes aquatic and terrestrial inverte-
brates but is rather varied. Among the more common prey
are caterpillars, crickets, locusts, beetles and their larvae.
Compared to the Australian White Ibis, they take more
small insects and spiders and fewer crayfish. In the dry
season, beetles, spiders and small lizards particularly are
taken. As noted above, the Strawnecked Ibis eats plague
locusts; Cowling (1974) found it prefers live to dead locusts,
but is nevertheless attracted to those rendered sluggish by
pesticides.
While nesting in suitable freshwater wetlands in the rainy

season, Strawnecked Ibises forage on flooded areas where
food is temporarily available. There they feed on dormant
aquatic prey and terrestrial animals drowned by rising
water levels (Carrick 1962). Continued flooding increases
plant growth, which supplements the supply of herbivorous
insects.
In most seasons, a bird will spend much of its time
foraging, occupying more than 75% of its time budget
(McKilligan 1975). Feeding activity tends to cease in mid-
day. Travel time is minimized by choosing roost sites, often
in woods or tall eucalyptus trees, near feeding and drinking
sites.
BREEDING
During its nesting season, the Strawnecked Ibis moves to
areas that flood seasonally and there takes advantage of
temporally abundant food supplies (Carrick 1962, Water-
man et al. 1971, Gosper et al. 1983). Because of the depen-
dence of breeding on the rainy season, it nests most reliably
in areas where water supplies are dependably seasonal.
McKilligan (1975) stated that breeding in southwestern
Australia is well defined from August to December. In
South Australia and Victoria much the same pattern
emerges. Occasional breeding takes place in the Northern
Territories from January to March, though this is on a very
small scale. In New South Wales and southwestern Queens-
land, the breeding season is not well defined (Carrick 1962).
We did not find evidence of breeding in colonies containing
Australian White Ibis on the Adelaide and Alligator Rivers,
N.T., even though rains are reliable in this area.
Nesting is an annual event in southeastern and south-
western Australia (Victoria, New South Wales, Western
Australia) but may not occur each year in the Northern
Territory or in Queensland, where nesting occurs in isolated
breeding colonies. Although usually absent from the arid
interior (McKilligan 1975), the species has been known to
nest there after heavy rains (Waterman et al. 1971).
Strawnecked Ibises begin to nest several months after the
beginning of the wet season (Carrick 1962, McKilligan
1975), unlike the Australian White Ibis which initiates
nesting soon after the rainy season begins. In some cases,
nesting is begun too late, the flood subsides and nesting is
abandoned.
Regional differences in the timing of nesting over Austra-
lia are explainable by variability of the timing of the rainy
season, and the situation is similar to that described for
Australian waterfowl (Braithwaite and Frith 1969). In per-
manently flooded swamps, temperature may also be a factor
in determining nesting season (McKilligan 1975).
Nests are built in low bushes, or on well-protected
reed beds on islands. Sometimes, however, the Strawnecked
Ibis will nest on dry ground where there are no trees,
bushes or reeds (Carrick 1962). Birds have been reported as
nesting in flood debris and large lignum clumps (Waterman
et al. 1971). They will roost in, but apparently not nest in,
trees.
Breeding is in large colonies, although sometimes only a
few pairs are to be found in mixed-species colonies of
Strawnecked Ibis 229
Australian White Ibis and herons. Sometimes they will take
over the breeding sites of the earlier-nesting Australian
White Ibis after the latter has raised its young.
Nothing has been recorded of the display of this species,
nor of its behaviour at the nest.
The nest is built of sticks, reeds and rushes. It is a shallow
cup, usually sited in a bush over water. When nests are in
reeds, the surrounding vegetation is trampled down, and the
areas becomes a single platform of vegetation.
Clutches vary from 1 to 4. Lowe found that in one year
the majority of the nests contained 2 eggs, with a mean of
2.29 eggs per nest, whereas the following year the mean
clutch was 2.42. Eggs are dull white but soon become
stained brown (Pizzey 1980).
Incubation takes 3i weeks, and fledging occurs 4 weeks
after hatching (Carrick 1962). Both parents feed and brood
the young (Carrick 1962). Feeding is by regurgitation and
continues up to 2 weeks after the young have left the nest.
Predation is not great, but food availability determines
success of the number of young raised, and when its supply
ceases young are readily abandoned. Lowe found that in one
year 112 clutches produced 52 fledged young.
TAXONOMY
Although long considered to represent a distinct genus (Car-
phibis), the Strawnecked Ibis appears to be fairly closely
related to the Sacred Ibis group (Holyoak 1970), one of
which—the Australian White Ibis—is also found in Austra-
lia. It differs primarily in its dark plumage, ornamental neck
plumes, and lack of ornamental display plumes derived
from the secondary feathers. It also occupies a slightly
different ecological niche, being better adapted to feed in
dry conditions and nesting on a schedule slightly delayed
behind that of the more aquatic Australian White Ibis.
A subspecies fozroi was described by Matthews based on
its being a smaller size than other populations. However,
the principal correlate of size differences in this species is
sexual, not geographic (Amadon and Woolfenden 1952).
CONSERVATION
Historically, the Strawnecked Ibis played an important role
in the lives of aborigines, who used it as part of their natural
diet. European occupation and habitat changes have ben-
efited the Strawnecked Ibis through the clearing of forest
and woodland, and the drainage of swamps for pasture and
farm land (McKilligan 1975). The bird is welcomed in dry
farming areas where it is recognized as a destroyer of insect
hordes. There seems to be no threat to the Strawnecked Ibis
in its dry-season refuge areas.
However, the Strawnecked Ibis does depend on fresh-
water swamps for nesting. Further wetland drainage and
flood prevention will restrict the available nesting areas.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 319.
 Waldrapp Ibis
Geronticus eremita (Linnaeus)

Upupa Eremita Linnaeus, 1758, Syst. Nat., ed. 10, p. 118: Switzerland

Other names:
Hermit Ibis, Bald Ibis, Bare-faced Ibis, Red-cheeked Ibis, Northern Bald Ibis, Crested Ibis, Red-
faced Ibis (English); Heremietibis (Dutch); Ibis chauve (French); Ibis calvo (Spanish); Waldrapp or
Waldrappe is used in German and Swiss as well as English

IDENTIFICATION
The Waldrapp is a dark, relatively large ibis, 70-80 cm long, with a bare red head set off by an untidy
ruff of feathers. Bannerman and Bannerman (1968) describing the striking plumage called the Wal-
drapp a grotesque, yet handsome bird.
The base plumage colour of the adult is a dark metallic green, which is tinged with a purple
iridescence. Its upper wings are glossed with copper and violet, forming a wing-patch that seems more
maroon in specimens we have seen. Elongated purple-glossed black feathers form a loose ruff or collar
emanating from the back and sides of the neck. The neck tuft, which is used in displays, gives this bird
its unkempt, dishevelled look.
The head is featherless, and the face and throat are red, except for the black crown topped by a
central streak of orange-red, which varies with age. The iris is grey, and the curved bill, along with the
legs and feet, are a dull red.
The sexes have similar plumage, but the male is slightly larger, a difference that is discernible among
groups of wild birds. Sample measurements are: wing, 40.3-42.0 cm (male) and 39.0-40.8 cm (female);
bill, 13.3-14.6 cm (male) and 11.5-13.1 cm (female). Siegfried (1972) found less sexual difference in bill
length in Moroccan birds than in eastern birds. In both sexes, the red of the soft parts brightens during
courtship.
Nestlings are covered with a grey-brown down, darker above than below and have a pink and black
bill. The juvenile is similar to the adult, but duller with very little iridescence and no copper coloration
on the wing. Unlike the adult, the head of a young bird is feathered (grey with white edges). The ruff
feathers are also underdeveloped. This permits recognition of young in the field without difficulty. The
head remains feathered through the second year. The eye is yellow-grey to grey, turning red only in the
bird's third year.
Generally, this is a silent bird when foraging, but noisy at the roost and at breeding sites. It uses bill
clappering at the nest, and its calls are either guttural or high pitched. The nearest description is that of
a domestic turkey. Such noises are made when bowing to Greet a mate, by an unpaired male at a nest
site, and at the time of nest-relief. Young give a call described by some as 'Lib, lib' when food Begging
(Brown et al. 1982).
The Waldrapp's flight is like that of a gull, taking a few wing-beats followed by a glide (Bannerman
and Bannerman 1968). Birds fly to and from roosts in V-formation, using shallow wing-beats inter-
spersed with glides on slightly bowed wings. In flight, the short legs are hidden beneath the tail.
The Waldrapp Ibis is a rather distinctive bird within its much restricted range. It is larger and
heavier than the Glossy Ibis, having shorter legs and neck. It is also distinguishable by its head and bill
colour.
232 Waldrapp Ibis
DISTRIBUTION AND POPULATION
Historically, the Waldrapp Ibis nested in the mountains of
central Europe, across northern Africa and into the Middle
East. But this range is now much reduced. Nesting is now
confined to Morocco, irregularly in Boghari in Algeria, and
in Birecik, Turkey.
The Waldrapp occurred in central Europe from the 12th
to the 17th centuries, ranging from southern Germany and
Austria in the valleys of the upper Rhine and Danube
Rivers, and in the Alps of Switzerland, Italy and Germany,
and perhaps in Hungary and Greece (Rothschild et al. 1897,
DesFayes 1987). In Africa and the Middle East, it occurred
in two distinct populations. The first nested in Morocco and
sporadically in northern Algeria; the second nested in south-
eastern Turkey and Syria (Geroudet 1965a, Kumerloeve
1978a, 1984, Cramp 1977, DesFayes 1987).
Evidence is lacking for reported nesting elsewhere, in-
cluding Yemen, Iraq and Ethiopia (K.D. Smith 1970). It
remains possible that it bred in ancient times in the Red Sea
Hills of Egypt and Sudan, where it was depicted as early as
2430-2410 B.C. and was part of the symbolism of ancient
Egypt (Meinertzhagen 1930, Kumerloeve 1983).
Migratory patterns differ between the eastern and west-
ern populations. Certainly, the population that previously
nested in central Europe would have migrated south in the
winter, and the eastern-nesting birds are still migratory.
These birds, nesting in Turkey, winter in Ethiopia, espe-
cially along the Red Sea (Bourne 1959).
The Waldrapp Ibises nesting in the west may only experi-
ence a post-breeding dispersal rather than a true migration
(K.D. Smith 1970). The birds nesting in Morocco disperse
from their colonies in June and July, and return in February
and March. Their wintering grounds are poorly known, but
many individuals have been reported remaining in Morocco
and in nearby Mauritania. It is estimated that 70% of the
population remained in Morocco for the winter, using three
main areas including the Sous and Massa Estuaries (Hirsch
1979b, Thevenot et al. 1982). Even in the middle Atlas
Mountains, birds appear to remain all winter (Snow 1952,
Hirsch 1982b).
Most immature birds do not return to the colony site
during the nesting season, and, overall, the Waldrapp Ibis
appears to be rather nomadic. It has been reported casually
in Iraq, Israel, Saudi Arabia, the Nile Valley of Egypt and
Sudan, Yemen, Somalia, Mali, Tunisia and Spain, and even
as far as the Cape Verde Islands and the Azores (Hartert
and Ogilvie Grant 1905, Salvadori 1899, Valverde 1959,
Bannerman and Bannerman 1968). The source of some of
these nomadic birds is not known for certain, leading to
speculation that undescribed colonies exist in Turkey or
elsewhere (K.D. Smith 1970), though the possibility seems
to be increasingly remote (Hirsch 1979a).
The world's population of this species is small, and con-
tinues to decrease, and as a result the Waldrapp is severely
endangered (Pegoraro 1989, Department of the Interior
1990). In 1982, the known breeding population was about
550 birds in 13 colonies, 12 of which were in Morocco, and
an additional 408 birds in 33 zoos (Michelmore and Oliver
1982). By 1989, there were about 750 birds in captivity. The
several nesting populations have their independent numeri-
cal trends.
The Moroccan subpopulation, nesting in many small
colonies and some isolated nests, has been decreasing
gradually for decades. At the beginning of the century, the
Waldrapp Ibis was an abundant species in Morocco, with a
large number of colonies being described by many authors.
Meade-Waldo (1903) described a colony on the cliffs of
Sale, with each bay occupied by many birds, so thick that
the nests touched. By 1920, however, this colony had been
extirpated (Jourdain 1922, Bede 1926). The population
stood at 1500 birds in 1940 (P. Jackson 1978). In 1965,
Geroudet (1965a,b) reported that the species was in "alarm-
ing decline'. By 1977, only 13 colonies were extant (Hirsch
1976b). In 1982, the total population there was estimated to
be 350-400 birds (367 were counted), including 93 pairs in
12 colonies (Michelmore and Oliver 1982). In 1982, Hirsch
(1982b) found 5 colonies in the High Atlas and adjacent
plains in eastern Morocco, containing 26 pairs and a total of
108 birds. Two colonies on the coast, outside the proposed
Massa National Park, contained 10 pairs, with a total of 76
birds. 5 colonies inside the proposed park contained 57 pairs
and a total of 183 birds. By 1984, this had increased to 220
birds. In May, 1989, only 43 pairs were counted in the
coastal colonies (Pegoraro 1989). However, in 1990, 78 pairs
fledged 72 chicks, and the Morocco population of about 300
birds is now considered reasonably stable (U. Hirsch pers.
comm. 1990).
The subpopulation nesting in nearby Algeria has always
been erratic, but continued to breed there periodically into
the 1980s. A small colony was discovered in 1974 in the

Djebel Amour near El Bayadh in the central Saharan Atlas
Mountains (Ledant et al. 1981). It consisted of 8-12 pairs
each year, and never more than 14 (Hirsch 1981b,d, 1982b).
In 1982, 14 nests fledged 10 young (R.E. Johnson pers.
comm. 1990). By 1984, only 7 birds were seen. By 1990,
about 5-12 birds arrived annually but did not breed because
of drought. (U. Hirsch pers. comm. 1990). A second colony
at Kasar el Boukhari had only up to 12 pairs and appeared
to depend on rainfall, which, when absent, resulted in this
nesting site being left unoccupied. Sporadic breeding may
have occurred up to 1953 at Kasar, where recently used
nests were discovered in that year (Kumerloeve 1958). The
colony site is now considered to be abandoned (Ledant et al.
1981).
The colony in Birecik, Turkey, was for many years the
stronghold of the eastern population segment. Some 3000
pairs nested there in 1890 (Kumerloeve 1962). Weigold
(1912-13) estimated 1000 birds there in 1912. This site
supported 1300 birds in 1953 (including immature birds
that were up to one-third of the total), 200 pairs in 1961, and
5 pairs in 1984 (Kumerloeve 1962, 1965, 1984, P.Jackson
1978, Hirsch 1982b, Luthin 1984a). By 1989, only 3 indi-
viduals returned to the Birecik colony, and 2 of these birds
were killed in a storm. So the colony now appears doomed
(Baris 1989a, Pegoraro 1989, H. Peters 1990).
These population decreases have been occurring for a
long time, with each population trend appearing to be
rather independent. It seems worthwhile to consider in
more detail the historic and recent status in segments of its
historic range, other than those where it has nested most
recently.
Any nesting that had occurred in Egypt ended millennia
ago. European nesting sites were last occupied in the 17th
century. In Syria, the last breeding was in 1916 or earlier. In
Iraq, there have been no records since the 1920s (Allouse
1953, Cramp 1977), and there is no proof that the species
ever bred there, contrary to Bannerman and Bannerman
(1968).
In Israel, there are no breeding records. A few records of
non-breeding birds have occurred since 1935. Up to 16 birds
were seen at Elat and one in 1980 near the Dead Sea (Hirsch
1980a, Lambert and Grimmett 1983). Attempts to release
captive birds into the wild have been unsuccessful. We
witnessed these birds from the Tel Aviv University flying
free around the area, perching on the balconies on nearby
apartment buildings.
In North Yemen, birds have been seen in marshes near
Ta'izz. 14 birds, including 2 juveniles, were seen in autumn
1985. In 1986, 9 birds were seen roosting with Cattle Egrets
(Bubulcus ibis) in January, and in January of 1987 at least 4
birds were seen again at Ta'izz. There is speculation as to
whether these birds might come from Turkey, or from some
undiscovered breeding site (Hirsch 1977a, Brooks et al.
1987, Bezzel and Wartmann 1990).
In Mali, single birds have been seen on the northern edge
of the Niger inundation zone, but certainly not in the num-
bers suggested by K.D. Smith (1970). In Mauritania, how-
ever, a flock of up to 50 birds was seen in February 1951 at
Bir Magrein in the northwest of the country, and about 20
Waldrapplbis 233
birds were seen in the south near Avatane Wells in the Djouf
(Dekeyser 1954, Valverde 1957, K.D. Smith 1970).
ECOLOGY
The Waldrapp Ibis is a terrestrial ibis, but both present and
past distributional records suggest that it is not confined to
such habitats (Meinertzhagen 1940, H.B. Peters 1928,
Hirsch 1981b). Typically it feeds in mountain meadows,
stubble fields, short grass, rocky semi-arid ground, on damp
ground in lower areas of high plateaus. The colony site at
Birecik is on and near the banks of the Euphrates River, and
those in North Africa are along streams or on the coast, so
they also feed in wetlands, on coastal sandy strips, on river
beds and in sandy banks on the seashore. They may also be
found feeding amongst cattle and village huts. Ruthke
(1977) observed them feeding at a rubbish dump. They
avoid tilled fields and pastures, especially if the grass ex-
ceeds c. 25 cm.
Despite its extensive use of terrestrial habitats, the Wal-
drapp Ibis appears to be dependent on rainfall, which
probably determines the timing and extent of the nesting
season and its nomadic movements (K.D. Smith 1970). In
winter, birds in the Ethiopian highlands feed on high moors,
wet meadows, and by fast-flowing mountain streams and
lake margins, up to 3500 m above sea level (Ash and Howell
1977).
Birds feed singly or in loose parties, by Pecking and by
Probing deeply in the ground, beneath stones, or under
plants and clumps of grass. Characteristically they feed
while Walking rather rapidly and will Run after insects. The
prey is lifted by the tip of the bill and quickly swallowed.
Hirsch (1979b) stated that these birds seldom drink, obtain-
ing most of their liquid intake from their food. In fact we
have not found any observation of how they catch fish in
water. They feed all day long.
The diet is broad, reflecting their ability to exploit what-
ever local prey abundances might occur (Meinertzhagen
1954, Siegfried 1966a, K.D. Smith 1970, Pala 1971, Parslow
1973, Porter 1973, Rencurel 1974). Invertebrates eaten in-
clude locusts, crickets, grasshoppers, ants, beetles, spiders,
caterpillars, earwigs, scorpions, centipedes, wood lice,
earthworms and snails. Vertebrates include fish, amphib-
ians, lizards and snakes, small rodents, and small birds
either live or dead, including nestlings. Vegetation eaten
includes berries, shoots, duckweed and rhizomes of aquatic
plants.
Favourite prey for birds in Moroccan colonies are snails
picked from shrubs, and in Birecik they eat larvae of beetles
and other insects. Aharoni saw one decapitate a young
horned viper (Cerates cerates). One Turkish bird contained
95% locusts (Parslow 1973), and one in winter from South
Yemen nothing but black beetles (Meinertzhagen 1954).
Hirsch and Schenker (1977) reported that the majority of
the food of birds at Birecik consisted of mole crickets (Gryllo-
talpa gryllotalpa), and various beetles and their larvae.
Roosts used away from the breeding area can be in trees,
on cliff ledges or sometimes in fields. On landing at the
roost, birds raise their plumage, croak and peck at each
234 Waldrapp Ibis
other. They interrupt feeding to sunbathe with well ex-
tended, drooping wings.
BREEDING
The Waldrapp Ibis nests in large or small colonies or alone.
The Moroccan and Syrian birds tend to nest in many small
colonies whereas the Turkish birds formerly nested in large
assemblages. Nests are placed on the ground, among
boulders on cliffs, on mountain ledges or in holes in rocky
cliff faces. These colony sites may be 100 m above the
ground. Nest-sites are usually near a river or next to the sea.
Historically, Waldrapps nested on castles in Europe and
have used artificial platforms erected on their behalf in the
Turkey colony site (Robin 1973).
Nesting occurs in March and April. However, in dry
years, most colonies on the plains are abandoned (Robin
1973).
The behaviour of the Waldrapp Ibis at its breeding sites
has been intensively studied, and the results have been
supplemented with extremely detailed accounts of behav-
iour under captive conditions (Mallet 1977, Hirsch 1979b,
W.L.R. Oliver et al. 1979, Thaler et al. 1981, Michelmore
and Oliver 1982, Sahin 1982a,b, 1983a,b,c, Pegoraro 1985).
The first birds to occupy breeding sites are males, who
acquire and defend a territory with fierce fighting. The
Forward Threat involves thrusting or touching with the
mantle held erect, the head retracted, and the bill raised and
gaping. Birds may beat their wings and grab and pull each
other backwards and forwards. Males stand at the potential
nest site calling. Activity increases with the arrival of the
females, who fly in circles to which the males respond with a
guttural 'Hoop hoop' call, while lowering their bill to touch
the breast, after which it is raised sharply. This is probably
the Bill Popping display. The approach of another bird
intensifies this action. The male crouches, passing its bill
under its flanks, simulating the insertion of nest material,
whilst swaying and making a rumbling sound. If the
approaching bird is a female, she will remain inactive until
approached but will then lower her head, soliciting allo-
preening of the neck, back and head feathers. During this
action the male continues to utter his 'Hoop, hoop' call.
Following allopreening, billing and entwining of necks takes
place; the male moving around the female. Such activities
may take place for up to a week before pair-formation is
completed.
Once paired, the arrival at the nest site of one of the birds
initiates a mutual Greeting display. Facing each other,
heads are first raised and then lowered so that the open bills
are nearly touching the ground. This is repeated two or
three times; on each occasion the crest and breast feathers
are raised and each bird utters a 'Jum' call as the bill is
lowered. At other times a slower more deliberate deep
bowing is followed by an upright posture, without a call.
Such ceremonies are formal and quite deliberate and are
followed by self-preening. They seldom occur away from the
nest site.
Male birds, after selecting a favoured site for nesting, will
clean away all remnants of previous nesting material and,
after the pair-bond has been formed, the male will collect
new material, arriving at the nest to present sticks to the
waiting female. Males will also steal sticks from nearby
nests. Both birds assist in building the nest. Nest-building
takes about a week to complete.
The nest is a loosely constructed structure of twigs and
sticks. The birds place a softer lining in the nest-cup, con-
sisting of leaves, grass and paper or other waste materials.
Promiscuity is reported in both captive and wild birds,
although, in observations of captive birds, approaches by
males to other females are usually rejected. Intrapair copu-
lations occur throughout the nesting period, long after in-
semination could occur (Sahin 1983a). This behaviour
appears to be a means of communication and reinforcement
of pair bonding in this species. Such behaviour especially
occurs in the event of breeding failure at the first attempt,
and sexual activity increases amongst unpaired or failed
pairs towards the end of the first nesting period (W.L.R.
Oliver et al. 1979).
Clutches are 3-A eggs. However, often only 2 are laid and
only 1 may be laid during droughts (Aharoni 1911, 1929).
Rencurel (1974) found that nests constructed in holes
usually contained 4 eggs whilst those on ledges only 3 or
even 2. The eggs are oval and rough pitted, their bluish
white colour becoming stained with brown during incu-
bation. Eggs are 63 X 44 mm, and the average weight is 68 g
(Schonwetter 1967). Double clutching can occur (Michel-
more and Oliver 1982, Sahin 1983a).
Eggs are laid 1 or 2 days apart (Hirsch 1979b, W.L.R.
Oliver et al. 1979). Time to hatching averages 28 days. Both
birds share incubation; and throughout incubation, the
incoming bird often presents nesting material and may have
to push its mate off the nest. Hatching success is low. Both
birds feed young by regurgitation, in captivity more feedings
are made by the male. Non-parental birds also feed young
(Sahin 1982a).
Eggs hatch 2-3 days apart, and sibling competition is
severe, even in captivity where food is readily available.
Overall nesting success is low, owing to hatching failure,
low food supplies, disturbance and, perhaps, pesticides. In
Morocco, 20% of the nests reared no young, 60% 1 young,
15% 2 young and only 5% 3 young (Cramp 1977). In dry
years, half the pairs reared no young and most of the rest
only one. Chick stealing by neighbouring pairs occurs some-
times.
Nestlings fledge after 43-47 days, and the young leave the
nesting site to travel to the feeding grounds with their
parents. Soon after the young join their parents in the
feeding areas, the colony site will be abandoned for the year
and nearby roosts occupied at night. The young eventually
separate into their own flocks (Ruthke 1977).
This ibis does not breed until its fifth year even though it
has acquired full adult plumage after 3 years. Older birds
appear to be more successful in rearing nestlings than do
younger birds.
TAXONOMY
Although considered to be monotypic, the eastern popu-
lation of the Waldrapp Ibis may have shorter bills than
 Waldrapplbis 235

western birds, leading Siegfried (1972) to suggest that they small Turkish town of Birecik has been the location of much
were isolated. of this activity. Hans Kumerloeve, followed by Udo Hirsch,
made valiant efforts to document and save the remaining
CONSERVATION birds at this colony (Kumerloeve 1962, 1967, 1984, Hirsch
1976b, 1978b). Studies, publications, appeals to inter-
The Waldrapp Ibis is extremely endangered, owing to long-
national organizations, lectures and the assistance of local
term population declines, which have accelerated in recent
politicians created some support. Locally a Waldrapp Ibis
years. In 1990, it was proposed for inclusion in the Endan-
day was initiated. In 1973, the World Wildlife Fund sup-
gered Species List by the USA (Department of the Interior
plied Hirsch with funds to increase his attempts to support
1990). This relict species has attracted an enormous amount
the colony. Wooden platforms were built; and, with diffi-
of attention from scientists and conservationists. In spite of
culty, birds were encouraged to use these safer nesting sites.
these considerable efforts, it is likely that the bird may
Nets were secured below the platforms to save young who
become extinct in the wild very shortly. Although all of the
fell out of the nests due to disturbance and an attempt was
reasons for the population decrease and localized extinc-
made to hand-rear some of these. In 1983, the returning
tions are not understood, several causes are identifiable.
adults abandoned the traditional colony site to nest near the
Over the past several thousands of years, climatic change
aviary holding captive nesting birds. This placed the wild
has probably led to a gradual range contraction. The species
birds in a safer situation.
occurs in Pleistocene fossils in Europe, and the climate there
In Morocco, disturbance by local people, tourists, and
prior to the 16th century was different from that of today.
egg and zoo collectors has similarly reduced the colonies,
Similarly, the climate of Egypt was more conducive to the
and more protection is vital. Efforts are underway to protect
species 5000 years before present than it is now (K.D. Smith
the wetlands critical to the population, at the Souss-Massa
1970). Over the shorter term, rainfall variations have coin-
Estuary (Salathe 1990).
cided with fluctuations in the colonies in Asia Minor and
Pesticide use has been and may continue to be one of the
Syria (Safriel 1980). However, failure of the population to
most severe threats to the species, which appears to be
respond to more recent climatic amelioration suggests that
extremely susceptible (Parslow 1973, Department of the
factors other than climate are now holding numbers down.
Interior 1990). Dry-land pesticides may have decimated the
Waldrapp Ibises were killed and eaten in Europe and
Turkish population, and residual poisons in fish stocks in
Africa, according to early accounts going back to the 1500s
both fresh and marine habitats may well have further affec-
(Jourdain 1934, Hirsch 1977a). Extirpation of the popu-
ted already weakened birds on their wintering grounds.
lation nesting in Syria is particularly ascribed to hunting.
In zoos, breeding of captive birds is meeting with con-
Syrians had long believed that eating the flesh of this bird
siderable success. The Basel Zoo pioneered this work, and
transferred sacred powers from the bird to the eater, and so
strong success has been attained in Jersey, Tel Aviv, Inns-
the species was eliminated from Syria early in this century.
bruck, Nuremberg and Wuppertal. We have observed the
Conversely, the Turkish colony, known to have existed since
captive birds at the Jersey Wildlife Preservation trust and
1879, has been protected by the goodwill of the local human
the Tel Aviv University Zoo, where release programmes are
population, who traditionally identify them with birds re-
in effect. At Alpenzoo Innsbruck-Tyrol 150 young have
leased from the Ark by Noah. The decline of this colony
been produced since 1963 (Pegoraro 1989). Because as
began with local human population increases and changed
many as 750 Waldrapp Ibises are in captivity (all derived
agricultural practices. The nearby town has greatly
from Moroccan birds), the reintroduction of birds back to
expanded, its population nearly doubling with the immi-
their natural habitat is, perhaps, the species' best hope for
gration of Kurds and Syrian Arabs from about 1958. Thus
the future (Pegoraro 1989, Akcakaya 1990, K. Brouwer and
the colony was confronted by people with little history of
M.C. Coulter 1990). Attempts to release captive birds are
toleration for the colonial birds nesting in their midst, and
being made in Israel and Spain. Additional studies in prep-
disturbance was severe. Stone-throwing by children caused
aration for expanded release programmes are now needed to
panic and desertion. At about the same time pesticides
assess hazards and the benefits. In June 1991, a meeting was
against malaria and locusts came into use, and it is strongly
held to begin developing a conservation strategy for the
suspected that this contributed to the rapid population
Waldrapp Ibis, especially to manage a viable captive popu-
decrease at about 1959-60, a period when over 600 birds
lation for reintroduction programmes (K. Brouwer 1991).
were found dead in the area (Parslow 1973, Hirsch 1976b,
1980a).
This highly endangered species has attracted consider-
able attention from a wide range of observers and conserva- Note: Body measurements, egg measurements and nesting
tionists who have initiated attempts to protect breeding season data for this species can be found in the Appendix on
areas and the dwindling numbers of breeding birds. The page 319.
 Bald Ibis
Geronticus calvus (Boddaert)

Tantalus calvus Boddaert, 1783, Table Planches Enlum., p. 52; based on 'Gourly a tete nue, du Gap de
bonne-Esperance' of Daubenton, 1765-81, Planches Enlum., pi. 867: Gape of Good Hope

Other names:

Southern Bald Ibis, Bare-headed Ibis, Wild Turkey (English); Glattnackenrapp (German); Wilde-
Kalkoen (Afrikaans); Umowanele (Zulu)

IDENTIFICATION
The Bald Ibis is a strikingly iridescent dark ibis of relatively large size, about 81 cm long. It is brightly
coloured with a whitish head, red bill and fluffy neck-ruff.
The adult's base plumage colour is dark green, highlighted by iridescent green, bronze and violet
overtones and iridescent copper patches on its shoulders. The neck-ruff is formed by long, fluffy, hackle-
shaped feathers of a metallic blue-green hue. The ruff surrounds the whitish-tan skin of the bare head
and neck. The crown of head is bright red and domed posteriorly. The relatively short bill as well as the
feet and legs are dull red. The eyes are red to an orange red.
The bare skin of the crown turns bright red during the courtship period. The sexes are alike, although
the males are larger than the females, with longer bills and tarsi.
Nestlings are covered with a dark grey down, darker on the crown. The soft parts are also dark: the
bill is black with a grey tip; the legs and feet are blackish grey. The eye is dark blue.
Immature birds are dark iridescent green and lack the copper wing-patches. The head and neck are
feathered light grey. The crown lacks the red colour of the adult, and the bill is flesh-coloured with a
greyish base. At 1 year, the plumage is similar to the adult, but the copper wing markings are less
bright. There is some white on the face at the base of the bill. The crown becomes pinkish orange. At
2 years, the crown becomes wrinkly red and less pronounced in colour than the adult. The bare face is
white with some red spots, and the neck feathers begin to form the ruff.
This is a relatively quiet bird, having a feeble gobbling sound (leading to an old Afrikaans common
name of Wilde-Kalkoen or wild turkey—Elliot 1877). It is, however, noisy at the nest. In flight it will
utter a high-pitched call similar to a gull or a feeble edition of an African Fish Eagle (Haliaeetus vocifer).
It is described as a high-pitched 'Keeuw-klaup, klaup' by J. Vincent and G. Symons (1948). It also
sometimes calls when evading a predator, or when a flock gathers. At the nest it has a wide variety of
calls. A piping call is uttered with the bill pointing downwards or resting on the back. The tail is flicked
with each note. Similar pipings and gruntings are giver! when fighting and during courtship. Bill
clappering occurs before and after copulation. The young utter a shrill Begging call at the approach of
the parents.
The Bald Ibis has powerful wing-beats in flight and is very crow-like in action (J. Vincent and G.
Symons 1948). It flies by gliding and soaring, using air currents and flapping flight to gain altitude. It
descends in a long glide, with its bill pointed forward.
Within its range, the red cap, pale neck and wing-patch of the Bald Ibis are distinctive among ibises.
Contrasted with the Hadada Ibis, it has a shorter bill, appearing straighter, and in flight its wings are
more pointed.
238 Bald Ibis
DISTRIBUTION AND POPULATION
The Bald Ibis occurs in a restricted area of the high grass-
lands of South Africa, Natal (Transvaal, Orange Free
State), Lesotho, and Swaziland (Manry 1985a, Clancey
1964a, Siegfried 1971). It is no longer found in the Cape
Province, having disappeared during the present century.
Generally, this is a sedentary species, often remaining
near its nesting areas year round. However, some seasonal
movements do take place, as some birds move out of their
nesting area, but this shift is generally not more than a few
kilometres. Historically, some birds moved closer to the
coast in winter (Elliot 1877). It has wandered as far as Cape
Town, where a single bird was collected at Milnerton in
1906 (L.E. Taylor 1909), and also to Tigerhoek near Cale-
don and Little Namaqualand, at the mouth of the Orange
River (Siegfried 1966a, 1971, Brooke in press).
The species is rare. Until recently the population was
considered to be about 2000 birds, including non-breeders
(Siegfried 1966b, 1971, W.B. King 1981). Recent surveys,
however, estimate the population to be much higher. The
total breeding population is now estimated at 5000-8000
birds (Manry 1985a,b, Collar and Stuart 1985). The popu-
lation appears to be stable, although continuing to be threa-
tened (Jaarsveld 1979, 1980, Bonde 1981, Manry 1985a,
Luthin pers. comm. 1990),
The population is centred primarily in the Orange Free
State where 64 colony sites were known by 1980 and 2400
birds, including non-breeders, were counted in one season
(Manry 1985a, Luthin pers. comm. 1982). In Transva'ai in
1982, 492 pairs were counted at 44 colony sites, 2 other sites
not being censused. In addition, 1294 birds were counted at
non-breeding roosts (Allan 1982). In Natal, 50 colonies with
about 400 breeding pairs were known in 1982. In Lesotho,
Jacot-Guillarmond (1936) reported that the population was
decreasing in the lowlands, but it is still common in the
mountains. A small population in 2 breeding sites remain in
the western mountains of Swaziland (Manry 1985a). At
least 4 nesting locations are known in Lesotho.
Surveys to determine the population status of this species
are rather difficult, as it nests solitarily in some places. The
breeding season is staggered, and nesting can be sporadic
depending on rainfall conditions (Siegfried 1971). In Natal,
one survey noted a large decline at 1 colony but increases at
2 others (K.H. Cooper and K.Z. Edwards 1969). There are
undoubtedly single breeding pairs that have been over-
looked.
ECOLOGY
This is a dry-land ibis, preferring short grasslands and
cultivated pastures, resorting to water only for drinking or
bathing. In western Natal, 41% of the ibis* feeding habitat
use was in short grassland, 40% was in pasture, about 1 %
was on sport fields and lawns, and the rest was in active and
abandoned cropland and ploughed fields (Manry 1982).
Habitat use changes seasonally, grassland being used
exclusively in January, which is early in the wet season. The
birds then move to rye pasture in mid-year where they
remain until December. They avoid medium to long grass
and other vegetated areas. So, as the grasses grow too tall
for foraging, the ibises move to heavily grazed pasture and
rye fields. In the same way, they are attracted to native and
artificial grasslands and rye grass pastures. They are par-
ticularly attracted to burnt grasslands in the first week after
burning and then return again to forage in the new growth
following the rains (Manry 1984a). Arthropods are particu-
larly abundant in winter-burnt grasslands, and the ibises
feed on insects disturbed or killed by burning. Their return
to the site occurs after the insects have recolonized the
regenerating grass.
Natural lightning fires, control burning and accidental
fires provide much of the short-grass habitat on which this
species depends. Its dependence on fire and grazing is
longstanding—the former range of the Bald Ibis coincided
with that of the black wildebeest (Connochaetes gnou), sugges-
ting they have similar environmental requirements (Sieg-
fried 1971).
Manry (1983) pointed out that cultivation has been prac-
tised in the species' range in Natal as early as 1600 to 1700
B.C. He proposed that the Bald Ibis was associated with this
patchily distributed foraging habitat and has continued
feeding in such locations until modern times. Today, grass-
lands are burned in winter to create firebreaks, and again in
the spring to enhance fresh grass growth; ibises benefit from
both of these customs (Manry 1985b).
The birds feed by walking while Pecking and Probing the
soil. On burnt ground, they Walk Quickly picking up living
and dead insects. Although wary, they forage around cattle.
They also turn over dung and take advantage of plagues of
locusts and grasshoppers. They also feed quite close to
human habitation, as the local people are benign towards
them.
The Bald Ibis is extremely gregarious, feeding and roost-
ing in large flocks. The birds move rapidly over the ground,
well spaced out. Feeding flocks fly to a foraging area from
roosts. They often stay in feeding area for up to a week,
before moving to another food source nearby.
The diet is poorly known, but Bald Ibises are clearly both
hunters and scavengers. Arthropods (especially caterpillars,
beetles, grasshoppers), earthworms, snails, frogs, small

dead mammals and birds form the bulk of their food (Stark
and Sclater 1906, J. Vincent and G. Symons 1948, Milstein
1973, 1974, Manry 1984a). They readily take plague insects.
The Bald Ibis by no means disdains carrion and acts as an
efficient scavenger (Stark and Sclater 1906).
Milstein (1973, 1974) reported on Bald Ibises' habit of
collecting buttons, which he suggests they may mistake for
beetles. But he also found, at the nest, a hollow porcelain
tube, a beer-can pull-tab, a toothpaste cap and two metal
rims of vaccine bottle tops. In one small colony, he found
156 small buttons, items not readily available in the forag-
ing habitat.
At dusk the ibises assemble in their roosts, on cliffs or in
trees, in groups of 40-50 with other ibises and herons. Birds
fly to foraging areas from their roosts, usually located in or
near their breeding areas amongst rock cliffs. As many as
40-150 birds will share their roost with Cattle Egrets,
Sacred Ibises, and cormorants (Pocock and Uys 1967).
Many of the roosts have been in use for a very long time;
Pocock and Uys (1967) suggested that this might be up to
100 years.
Bald Ibises will sunbathe with wings wide open and
touching the ground.
BREEDING
This ibis is a ground-nester, especially on cliffs along deep
river gorges. It often nests by waterfalls, even in the spray
zone or under the falls. In other colonies, however, no water
is present (Milstein and Siegfried 1970).
Birds may nest alone, but the Bald Ibis is more commonly
a highly gregarious species nesting in colonies of 40 pairs or
more. Numbers vary at traditional sites from one year to the
next. Although sedentary for the most part, some interco-
lony movement has been documented (Milstein and Wolff
1973).
Claiming of potential nest-sites and pair-bonding occur as
early as June, as the birds spend increasing time at the
colony sites and less time off foraging. Males occupy the
nest-site first, defending them by bill jabbing and sparring.
Courtship behaviour has been difficult to observe. At the
display site, initial interactions seem perfunctory with copu-
lations appearing to be somewhat randomly directed at
different partners. A male and female will behave as a pair,
and then separate after a few days. A pair will move from
ledge to ledge, performing courtship displays. Mutual
preening takes place, particularly directed at the distinctive
neck feathers. Pairs grab each other's bills, wagging their
heads strongly. Twig presentation can be made by either
one of the pair, and this is accompanied by raising and
lowering of the sideways-turned head. A second bird will
often grab the twig and wiggle it.
After being apart for some time, the male will peck at the
female's bill, body, neck or wings with his closed bill. The
females responds by extending her neck vertically with the
bill resting against her neck, concealing her face and bill
within raised neck plumes. Sometimes the female will rest
the extended bill against her neck or on the ground.
To copulate, a male will bite a female's foot to induce
Bald Ibis 239
crouching, and then mount to copulate. During the brief
copulation, the male grasps the female's bill and shakes it
vigorously. Often the two birds will then stand erect with
bills raised, snapping their mandibles together.
Upon return of the mate during incubation or brooding, a
bird on the nest will flick its tail and make piping noises,
bending its neck so that the nape of its head touches its
back. Intruders receive snapping bills pointed at them.
Upon successfully repelling the invader, the bird on the nest
will rub faces with its mate. They squirm, twist their necks
and utter squealing noises.
The male collects nesting material for the female to use in
building the structure. The female presses her breast down
to form a hollow of the cavity (Brown et al. 1982, Manry
unpub. report). The nest itself is placed on a cliff ledge or in
holes in the cliff face. There is no confirmation of tree-
nesting, though the possibility is discussed by Pocock and
Uys (1967). The nest is quite large, about 50 cm in diameter
and 15 cm high.
Eggs are laid between August and October, in the dry
winter season. Egg-laying can be carried out over quite an
extended period. The clutch is 1-3 pale blue-brown spotted
eggs, averaging 1.9/nest in the Transvaal (Milstein and
Wolff 1973). Eggs are laid at 2-day intervals. They measure
63 X 43 mm and weigh about 68 g (Milstein and Wolff
1973, Milstein and Siegfried 1970). In Transvaal, 1.8 young
hatched per nest and in Natal 0.83, although this varies
according to the season (Milstein and Wolff 1973, Manry
unpub. report).
Both adults feed the young by regurgitation, which de-
velops into a tussle with older chicks, who Beg by head
bobbing while giving shrill cries. To provide food from its
gullet, the adult rests its bill on the floor of the nest. Older
chicks tap the bill of their parent after its arrival at the nest
and then insert their bills into the adult's opened mouth.
The young fly after about 55 days, in mid-October to
December (Manry 1985b), but depend on their parents for
food for up to 2 months.
Nesting success depends on available food supplies. In
droughts, many birds do not breed, and, when food short-
ages occur, the young starve (Manry 1985c). Ibises are also
lost to predation, including by Jackal Buzzards (Buteo rufofu-
cus), Cape Eagle Owls (Bubo capensis), Pied Crows (Corvus
albus) and man (Milstein 1973). Attacks by Black Eagles
(Aquila verreauxi) have been witnessed, and one pair of these
birds, which occupied the nearby cliffs, caused the Bald
Ibises to abandon their nests (Allan 1982). Overall, how-
ever, breeding success is generally good. In Transvaal,
South Africa, in 1983 about 400 pairs raised 277 young.
TAXONOMY
The genus Geronticus contains two species widely separated
in Africa. Both have similar plumage and nest on cliffs. The
southern species is sedentary, whereas the more northern
species is migratory and somewhat nomadic. The relation-
ship of the Geronticus to other ibises is obscure (Holyoak
1970), but Fry et al. (1985) suggested that Geronticus and
Bostrychia may be related.
240 Bald Ibis
CONSERVATION
The Bald Ibis has long played a role in human history
(Kumerloeve 1978a, J. Vincent and G. Symons 1948). His-
torically, young were taken to be used as medicine by the
Zulu. It certainly was once more widespread in southern
Africa than it is at present (Siegfried 1971, Milstein and
Wolff 1973). It is now confined to a limited range and man-
induced foraging habitat.
Hunting has always been a factor. The Boer farmers shot
this ibis during the South African wars to supplement their
meat supplies. Settlers also hunted it for both food and
feathers. Although the species is officially protected
throughout its range and colony sites are under protection
in Transvaal (Milstein and Wolff 1973), local hunting press-
ure remains, as people take young from the nest for food and
for other reasons. Such hunting may be the largest short-
term threat to the species. Other disturbance, as well as loss
of nesting sites, also threatens the bird.
However, the main long-term reason for the species' de-
cline during the present century was the spread of karoo
vegetation following severe overgrazing of the grasslands
(Siegfried 1966a,b, 1971, Brooke in press). The encroach-
ment of brush and karoo vegetation in the Cape Province
and afforestation by conifers in Natal and Transvaal cer-
tainly make the grassveld unsuitable (Siegfried 1971, Manry
1985a). This shy bird is now entirely dependent on farming
communities, who own the sheep and cattle pastures in
which they feed and the cliffs on which they nest. In the
Transvaal, cash incentives and warnings of prosecution
were successful in re-establishing one colony (Allan 1983a).
The habitat requirements of the species are very exacting.
For example, it is found only in areas where the grasslands
are burned annually, and not where the fire cycle is longer
(Manry 1985c). Some parts of its former range may be made
more suitable through restoration of its grassland environ-
ment, and irrigation schemes may be increasing habitat
quality and consistency.
A small captive population in the Pretoria Zoo may
become the core of a reintroduction programme (Brooke in
press). A determined programme of education, habitat im-
provement, and cooperation of land owners can save this
bird from further declines.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 320.
 Black Ibis
Pseudibis papillosa (Temminck)

Subspecies:
Pseudibis papillosa papillosa (Temminck)
Ibis papillosa Temminck, 1824. Planches Color., livr. 51, p. 304 and text: India and Ceylon (=Sri
Lanka)

Pseudibis papillosa davisoni (Hume)

Geronticus Davisoni, Hume, 1875, Stray Feathers, 3, p. 300: Pakchan Estuary, Tennasserim, India

Other names:
White-shouldered Ibis, White-necked Ibis, Red-naped Ibis, Burma Ibis (davisoni), Tenasserim Black
Ibis (English); Schwarzen-Ibisse (German); Ibis Noir Oriental (French); Baza, Kala Baza, Karan
Kul (Hindi); Kulo Dochara (Bengali); Kala Akohi Bog (Assam); Nella Kankanam (Telugu)

IDENTIFICATION
The Black Ibis is a dark ibis, about 68 cm long, distinguished by the white wing-patch that is visible
while the bird is in flight.
The common name belies the adult base colour of this ibis, which has a dark brown body, neck, head
and wings. The dark, glossy blue secondary feathers of the wing are sometimes flecked with white. A
large, broad patch of white runs along the leading edge of the wing on the lesser wing-coverts to well
below the carpal joint. Although prominent in flight, this feature shows only as a thin white line when
the wings are closed. The short legs are hidden beneath the steel-blue tail in flight. The tibia has
feathering halfway to the tarsus joint.
The head, somewhat square shaped, is black and .distinctively capped with a triangular patch of
warty red skin, absent in davisoni. The eye is orange-red to bright red. The curved bill and the broad,
stubby feet are horn coloured. The iris, legs and feet turn bright red at the commencement of breeding.
The male is slightly larger than the female.
The subspecies davisoni and papillosa arc distinguished by the former lacking the red cap.
Young birds are dark brown all over, with paler breasts. They lack the iridescent tinge on their
feathers. The pale horn-coloured bill is shorter than that of the adult.
The Black Ibis is generally a silent bird when feeding but is noisy at the evening roost. There, its call
is loud and trumpet-like. It is sounded particularly when the breeding season approaches, as the male
birds advertise at nest sites prior to pair-formation.
This species can be confused at a distance with the Glossy Ibis, but the Black Ibis is larger and
shorter legged and is clearly distinguished by the brilliant red head (in papillosa) and white wing-patch.
In the Mekong Delta, it is said to join feeding associations with the Giant Ibis, but the latter is
substantially larger. In all comparisons, the white wing-patches distinguish this species in flight.

DISTRIBUTION AND POPULATION

The Black Ibis was formerly widespread in the Indian subcontinent through Southeast Asia, in
Pakistan, Nepal, India, Burma, western China, Thailand, Laos, Cambodia, Vietnam and the Malay-
sian Peninsula.
The western subspecies is papillosa, formerly occurring in Pakistan and Nepal, south to India (Mysore
and east to Bengal and Assam), and Bangladesh.

The blue ? indicates the race davidsonii.
The eastern subspecies is davisoni, formerly found
throughout Southeast Asia, from Burma, to northwestern,
central and peninsular Thailand, Malaysia, Cambodia,
Laos, Vietnam (to southern Annam and Cochinchina), and
north into Yunnan, China. It has been reported from Bor-
neo for many years, but its breeding status has not been
documented.
The species is now rather patchily distributed throughout
its remaining range. In Pakistan, where it was once wide-
spread, occasional sightings are made during the rainy
season only. On the Indian sub-continent, it is found in
small populations (S. Ali and S.D. Ripley 1968, T. Roberts
pers. comm. 1990). It is less rare in Rajasthan southward
through India, and in Gujarat it is still quite common (D.A.
Scott 1989). In Bengal and Assam, it is locally common. In
the Nepalese terai, it is found in some numbers along the
river banks. There is a doubtful record ofpapillosa from the
Arakan, which probablyvrefers to davisoni. It is not recorded
from Sri Lanka or the Andaman Islands.
The subspecies davisoni is increasingly rare and has been
brought to near extinction (Bain and Humphrey 1980). In
Thailand there have been no records since 1937, except for
one unconfirmed report from the Thale Noi wetlands (W.B.
King 1981), nor are there recent reports from Burma or
Malaysia. The last record from China was from Yunnan in
1889. Vo Qoy and Le Due (pers. comm. 1990) consider that
the Black Ibis is still to be found in parts of Cambodia and
Laos, and also in southern Vietnam near the Cambodian
border (Wildash 1968, D.A. Scott 1989). Two birds were
seen in Dong Nai, Vietnam in 1991 (Eames 1991). Since
1979, it has been reported, without total certainty, several
times from Indonesian Borneo, most recently from the Kali-
mantan wetlands in 1983 and from the Barito Swamps in
1989 (MJ. Silvius and WJ.M. Verheugt pers. comm.
1990). Silvius and Verheugt report that a small number are
believed to survive along interior rivers in Kalimantan,
Indonesia.
Black Ibis 243
The species is generally non-migratory in India and else-
where. We have seen it in Gujarat even during droughts.
However there is some evidence for population movements
in response to monsoons, particularly dispersal of young
birds. Birds are found in Pakistan only occasionally and
during the rainy season. They also occur casually elsewhere
outside their normal range, such as in Burma and Borneo.
Population sizes are not known. Thailand was once the
centre of the subspecies davisoni population and it was once
common there (Bain and Humphrey 1980). In Gujarat,
India, V.C. Soni (pers. comm. 1990) of Saurastra Univer-
sity found 10 roosts totalling 325 birds in the Rajkot area
and a further 8 roosts totaling 270 individuals. We have seen
flocks of 20-30 birds in many parts of this state. S. Faisa
(pers. comm. 1990) found 30 birds in Tamil Nadu, where it
is generally rare, though we have seen them in small num-
bers in nearby regions. Other recent censuses include 200 in
winter at Chandpata Lake, Madhya Pradesh, and 45 in
winter at the Karnataka Tanks, Anantapur and Bangalore,
India. The Asian Midwinter Waterfowl Survey in 1988
recorded 220 in India (Rajasthan, Uttar Pradesh, Madhya
Pradesh) and 62 in Nepal (Royal Chitwan National Park)
(Van der Ven 1988). The population size of davisoni is
impossible to ascertain, if it survives at all.
ECOLOGY
The Black Ibis occupies a similar ecological niche in Asia as
does the Hadada Ibis in Africa. It prefers to feed near water,
but uses a wide range of aquatic and terrestrial foraging
sites, especially in shallow water, around the edge of
marshes, in other wetlands, jheels or man-made tanks, coas-
tal estuaries, river edges, paddy-fields, grasslands, culti-
vated fields. Where it is not persecuted, it will feed close to
human habitation, in urban areas foraging amongst open
sewage, water-well spills and flooded fields (V.L. Dharmak-
umarsnhji pers. comm. 1990, pers. obs. J.A.H.). Soni (1988)
identified several specific feeding sites, including crop fields
(usually after harvest), grassland, sewage, garbage, litter,
carrion, manure heaps, shallow muddy shorelines, estuaries
and marshlands.
Characteristics of behaviour varies, depending on the
history of disturbance. In India, the Black Ibis is not a shy
bird; it feeds almost under the feet of villagers, and when
disturbed it often flies round in a short circle to return to the
same area from which it took off. In Southeast Asia, how-
ever, where it is rare and widely persecuted, it is extremely
shy (Wildash 1968).
The Black Ibis is a tactile feeder, like all ibises, and
adapts the techniques used according to food taken. Its
feeding behaviours include Standing, Shallow and Deep
Probing, Pecking, Groping and Head Swinging. Soni (1988)
described the Black Ibis as using Groping and Head Swing-
ing along muddy shores. It will Stand quietly while Probing
deeply or shallowly, or Pecking.
In small parties, the birds Walk Slowly in the same
direction across a field keeping well spaced out and seldom
interacting. The birds remain separate from other species,
except when gathered at carrion and sewage, where they are
often in the company of Cattle Egrets (Bubulcus ibis), House
244 Black Ibis
Crows (Corvus spendens) and Jungle crows (Corvus macrorhyn-
chos). During pair-bonding, females will occasionally beg
food from males on the foraging grounds (pers. obs. J.A.H.).
Soni (1988) considered the simple repertoire of feeding
behaviour to be something of a constraint. The species is
nevertheless, rather adaptable. It can change diet according
to availability of food supply, as demonstrated during the
recent drought of several years duration in Gujarat. When
groundnuts ripen in Gujarat, huge flocks of Demoiselle
(Anthropoides virgo) and Common (Grus grus) Cranes feed
extensively on this food, and they are joined by Black Ibis.
With the lack of fresh water and failure of the crop, all the
cranes left the area, but the ibises remained, changing their
diet by exploiting the carcasses of cattle killed in the
drought. They fed on the carrion and on large quantities of
associated insect larvae. This food source, along with dom-
estic sewage, provided a major portion of the Black Ibises'
intake throughout the period.
The species' omnivorous diet includes earthworms, milli-
pedes, centipedes, larvae and pupae of house flies, rat-tailed
maggots, dung beetles, locusts, grasshoppers, thrips, scarab
beetles, cocktail beetles, white grubs, collembola, frog tad-
poles, garbage beetles, small crustaceans, lizards, small
slow-moving mammals, and even birds. It also appears to
consume groundnuts, and leaves of cultivated and wild
plants (Soni 1988, pers. obs. J.A.H.).
As dusk falls, the scattered birds collect to roost commu-
nally usually in large trees. Where there are no trees, we
have seen them use small islands or the centre of large fields,
where they will stand close together in the open and fly off
silently if disturbed.
BREEDING
In Gujarat, Black Ibis have two distinct breeding seasons,
one in October and November and a second more regular
one in March and April. These seasons precede the periods
of coastal monsoons. Elsewhere, the timing of the nesting
season varies geographically depending on the timing of the
monsoons.
The Black Ibis nests singly, usually high in a tree such as
a peepul or banyan, nearly always occupying an old nest of
a bird of prey, a vulture or a crow. Often the tree is very
close to human habitation. We have seen a nest used
annually in the grounds of the Palace of Jasdan, and in the
tall palm trees in the Palace grounds in Porbandar. Con-
versely we have seen several nests quite close together, with
at least 2 nests in one large tree in the riverine forest belt
along the rivers in the Nepalese terai.
Nests are often used annually in spite of nest failures due
to harassment by House and Jungle Crows. Courtship be-
haviour has not been well studied. Prior to pair-formation,
the male issues a trumpet-like call when advertising from its
nest site or on the wing. Copulation is on the nest or in a
nearby tree, but not on the ground. Both birds collect new
material for the nest and this activity continues well after
incubation commences.
Two to four eggs are laid. They are pale bluish green,
usually but not always spotted and flecked with red. The
average size of 59 eggs measured was 63.0 X 43.8 mm
(Baker in S. Ali and S.D. Ripley 1968). Incubation lasts
33 days (Kumar and Soni 1984). Young are fed at the nest
by regurgitation, and fledged young will still beg for food
after leaving the nest.
TAXONOMY
The two races of the Black Ibis were once placed in separate
genera (Inocotis papillosus and Graptocephalus davisoni). Elliot
(1877) and other authors recognized their similarity but
considered them to be seperate species, Pseudibis papillosa
and P. davisoni, and even today some authors give specific
status to each. Holyoak (1970) pointed out how similar they
really were, and despite their apparently allopatric distri-
bution, proposed that they be considered a single species,
which we and most others have followed.
CONSERVATION
The eastern subspecies of the Black Ibis (davisoni) is highly
endangered. In fact, this formally widespread species is
almost entirely dependent on human tolerance. Its adapt-
able food choice means that, without persecution, it can
survive under both wet and dry conditions in a wide range
of wet and dry habitat, including near habitation. However
without secure nest-sites, it has no chance of long-term
survival. Hunting in Pakistan, often for sport, and shooting
in Southeast Asia for food have played a large part in its
demise from those areas. In other areas, warfare has
reduced both populations and natural habitats.
It is a sad fact that former havens provided by religious
sites are now less available throughout the species' range.
The tolerance of the Hindu population has until now played
a major part in the Black Ibis' well-being in Gujarat,
although even here continuing drought and the felling of
trees present a growing threat to its survival.
A combination of the rapidly increasing human popu-
lation in all these regions, necessitating more intensive culti-
vation, draining of wetlands, use of pesticides, and nearly
continuous warfare has virtually wiped out this species, as it
has many other storks, ibises and spoonbills that were
formally present in very large numbers.
There is a need for increased surveys over the range of
this species (Holmes and Burton 1987), and for identifi-
cation and protection of nesting and feeding sites from
human intrusion. M.J. Silvius and WJ.M. Verheugt (pers.
comm. 1990) suggest that in Indonesia, surveys should be
conducted in the Mahakam Lakes region and upper
Mahakam River. The species must also be protected from
hunting throughout its range, as it is now in Thailand (W.B.
King 1978). The very recent discovery of two birds in
Vietnam, in the Nam Bai Cat Tien National Park, is a
matter of great importance. However, it is also a matter for
concern in that serious conflicts exist between park officials
and fishermen living in the park, which led to a shoot-out
and deaths (Anon 199Id). It is hoped that relocation of
residents will provide an opportunity to maintain these
wetlands.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 320.
 Giant Ibis
Thaumatibis gigantea (Oustalet)

Ibis gigantea Oustalet, 1877, Bull. Soc. Philomath. Paris, Ser. 7, 1, p. 25: Mekong River, Cambodia

Other name: Mekong Ibis (English)

IDENTIFICATION
The Giant Ibis is the largest member of the ibis family, up to 103 cm tall. It is a dark bird, with a
featherless black head and neck.
The base plumage colour of the adult is brownish. The lower part of the neck is covered with greenish
brown feathers edged with grey. The back is dark brown with a green gloss. The scapulars are dark
brown, the primaries brownish black, and the secondaries and wing-coverts dark green. The wings
extend to the end of the tail.
The head is large and nearly square. The head and upper neck are bare, covered with black skin with
prominent black bars across the head and back of the neck. The bill is brown to corn brown, or pale
brown (Delacour and Jabouille 1931, B.F. King and E.G. Dickinson 1975, pers. obs. J.A.H.). The legs
and feet are red to violet, and have also been reported to be straw brown (Delacour and Jabouille 1931,
Wildash 1968, B.F. King and E.G. Dickinson 1975, Lekagul and Cronin 1974).
For an ibis, this species is remarkably large. Example measurements are wing 52.3-57.0, tail 30.0,
tarsus 11.0, culmen 20.8-23.4 cm (Delacour and Jabouille 1931).
The sexes are alike, but females are, on average, slightly smaller than males. The differences in the
description of soft-part colours suggest that these change seasonally.
Little is known about the plumage of juveniles. One juvenile measured was smaller than an adult
(wing 53 cm and culmen 18 cm) (Delacour and Jabouille 1931). Elliot (1877) described one specimen
he considered to be a juvenile, but this description did not differ from that of an adult. It seems likely
that juveniles lack the iridescent gloss of the adult.
Its size alone will distinguish this species from other ibises. It has a heavy body, long neck and long
bill, though the tarsus is relatively short. It has been seen with the Black Ibis, which is smaller and has a
white wing-patch.

DISTRIBUTION AND POPULATION

This bird formerly nested in southeast Thailand, central and northern Cambodia, southern Laos, and
southern Vietnam (King in Collar and Andrew 1988). Its historic nesting range was centred in
Cambodia, extending into the Mekong Valley wetlands of eastern Thailand and the Saravane district of
Laos. It nested at Phu-Rieng near Saigon in 1925.
Fisher et al. (1969) suggested that it could be called the 'Mekong Ibis', to emphasize that its range
centres around this great river. In fact, the species has never been proven to breed outside of the
Southeast Asian mainland drained by this river.
The species is certainly extinct in Thailand (P. Round pers. comm. 1989). Fisher et al. (1969)
believed that its recent breeding grounds are scattered over a stretch of lowlands from the Mekong
River on the Laos border area, 325 km to the southwest into the valley of the River Sen for 50-65 km
north of Kompong Thorn. The latter region is where Delacour found the species numerous in 1927-28.

Vo Quy (pers. comm. 1989) considers that the birds remain
only in the Mekong Delta of southern Vietnam and along
the border of Cambodia. He suggests that they may be
found in the mangrove forests of Nam Can, the Melaleuca
forests of U Munh, and in Dong Thap Muoi. Indeed, C.S.
Luthin (in Collar and Andrew 1988) reported small num-
bers in southern Vietnam at Dong Thao Muoi, an inland
delta of the Mekong. Giant Ibises have not been seen in this
area, however, since 1980 (D.A. Scott 1989).
Birds have been collected as vagrants outside their usual
area. These were found in peninsular Thailand and Malay-
sia on the opposite side of the Gulf of Thailand (Fisher et al.
1969).
Little is known about population sizes. The Giant Ibis
was numerous in 1928 and 1929 in the valley of the River
Sen, north of Kompong Thorn (Deiacour and Jabouille
1931). Wildash (1968) found it quite common in places in
1968. Population estimates range from 100 birds (Bain and
Humphrey 1980) to statements that the species is extinct.
ECOLOGY
The Giant Ibis is a lowland bird, feeding around lakes, in
swamps and marshes, on open plains, in open forests and
clearings, and in ponds in deeper forests (B.F. King and
Giant Ibis 247
E.G. Dickinson 1975). One was collected from a paddy-field
in Trang in the Malay Peninsula in 1896. It is reported to
occupy the same feeding habitats as the smaller Black Ibis,
and to feed with it.
The birds appear to feed in pairs and small groups (Wil-
dash 1968, Milton in Fisher et al. 1969). In times of drought
they are said to aggregate at permanent water holes (Bain
and Humphrey 1980). Virtually nothing is known of their
behaviour. It is reported to be a shy bird.
BREEDING
Again, virtually nothing is known of the nesting habits of
this rare species.
TAXONOMY
Although for many years this distinctive, stork-sized ibis
was assigned to its own genus, some recent treatments
merge it into the genus Pseudibis, implying a close relation to
the Black Ibis (Pseudibis papillosa), with which it occurs (J.
Steinbacher 1979). Olson (1984) stated that the appropriate
anatomical study has not been done and cautioned against
relegating the species to Pseudibis without such a study. This
is a very distinctive ibis, and it is quite logical that this
distinctiveness should be recognized taxonomically until
morphological, behavioural and biochemical studies have
been completed that demonstrate otherwise. Therefore, we
have kept it in the genus Thaumatibis.
CONSERVATION
The Giant Ibis is in as much trouble as any of the endan-
gered ibises. It is not clear how many survive and where. As
the species account suggests, almost nothing is known of its
basic biology. This is a striking, unique ibis—owing to its
size, which is remarkable for a member of the family —and it
deserves to be saved from extinction. Unfortunately, no
conservation measures can be proposed because no study
has been made of the species in the wild. This must be
remedied: the present distribution must be determined and
secure reserves set up to accommodate the species.
Much of this species' habitat has been destroyed by
draining for rice and other crops and by the ravages of war,
which has plagued this region for decades. Add to this the
ever-increasing human population in parts of its range, and
it is clear that only protection in nature reserves can give
this vulnerable species a slim chance of survival.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 247.
 Oriental
Crested Ibis
Nipponia nippon (Temminck)

Ibis nippon Temminck, 1835, Planches Color., livr. 93, pi. 551: Japan

Other names:
Japanese Crested Ibis, Japanese Ibis, Rosy Ibis, Chinese Ibis, Asian Crested Ibis (English); Toki
(Japanese)

IDENTIFICATION

The Oriental Crested Ibis is a striking bird, with its rosy-tinged white plumage, dark bill and
pronounced crest. It is a medium-sized ibis, c. 56 cm long, appearing to have long wings and a rounded
tail in flight.
The non-breeding plumage is basically white. The crest feathers are sometimes tinged with pink, and
the back and wings are also tinted a pale shade of pink. We have seen museum specimens from Japan in
which the remaining coloration is orange. In flight, the underwings show quite strong pink colouring as
does the tail.
The black, downcurved bill has a red tip. The unfeathered portion of the face is a bright red. The iris
is red, and the eye has a yellow ring encircling it (G. Archibald and S. Lantis 1979). The legs and feet
are reddish brown.
Prior to the breeding season a gradual colour change takes place, as the head, back and neck take on a
grey coloration. This change is cosmetic, rather than structural, and originates in a black secretion from
an area of skin under the feathers surrounding the ibis' naked face. The bird applies this tint to its
feathers by daubing (Uchida 1970, 1974). The black substance adheres to the proximal feather barbules
creating a grey or blackish tinge, which becomes more marked each time the daubing takes place.
Firstly the crest, then the head and neck, and finally the back become dark grey. This coloration
remains throughout the breeding season until a post-breeding moult returns the feathers to white.
Immature birds have cheeks covered with downy feathers, and the rest of the face is a bare and
orange yellow in colour. The general immature plumage is a dusky cream, glossed with rose. The
primaries are a blackish brown, and the legs and feet light brown. The irides are a light yellowish
brown.
The Oriental Crested Ibis is generally silent. Its infrequent call is a 'Gak', given before taking flight
and while flying. It appears to be used as a low-intensity alarm call and for localization. A 'Gak-gak-
gak' is given in response to disturbance. A growl-like call is given in agonistic situations.
This is the only all-white crested ibis, and should not be confused with any other in its extraordinarily
limited range.

DISTRIBUTION AND POPULATION

Originally the core of the breeding range of the Oriental Crested Ibis was in eastern Asia. It occurred
from the Ussuri River in eastern USSR and in eastern Manchuria, as far north as Heilongjiang through
central China (south to Anhwei and Chekiang west to Shensi and Kansu), Korea, and east to Japan
(Dementiev and Gladkov 1951:433-437, Yamashina 1977a,b, Cheng 1976).
250 Oriental Crested Ibis
In recent years the bird has been known in the wild only
from Shaanxi Province, China, the Demilitarized Zone of
Korea, and Sado Island, Japan (G. Archibald and S. Lantis
1979).
At least the northern populations of China, Japan, and
Manchuria appear to have been migratory, moving in
autumn into southern Japan, the Yangtze River basin of
China, and the islands of Taiwan and Hainan. It is thought
that the recent sightings in Korea were of birds wintering
there. Southern populations in China and Japan were more
sedentary, remaining year-round except for localized move-
ments. In China, Yinzeng found that after fledging in July
the ibises moved to the lowlands 20 km from their nesting
area (W.W. Brehm-Fonds 1984).
Populations of the Oriental Crested Ibis were reasonably
large until the mid-1800s. They subsequently decreased due
to shooting, habitat loss, and pesticides. It is presently one
of the world's most endangered birds.
The changing population in Japan has reflected the level
of its protection. Before the Meiji Era, the species was
protected by the ruling classes and often seen in suburbs of
Edo (now Tokyo). However, from the Meiji Restoration in
1886 and the introduction of firearms, down to 1892 when
hunting restrictions were established for the first time, the
ibis and other large birds were hunted (Takashima 1957a).
The decrease was precipitous, occurring in the three
decades after hunting was permitted (G. Archibald and S.
Lantis 1979). The species apparently survived only in the
Niigata and Ishikawa Prefectures, on the Noto Peninsula
and on Sado Island, where it was rediscovered by Dr S.
Uchida in 1933 (Takashima 1957a). In 1936, 27 individuals
were known to exist, and for many decades this population
was the only known to survive. They last bred in the wild in
1973. In 1981 the remaining 5 wild birds were taken into
captivity, joining another bird, to form the nucleiis of a
captive breeding population. By 1984 the captive popu-
lation had fallen to 3.
The population in the USSR has never been well docu-
mented. It has not been observed there since 1983.
Although the present status there is unknown, on the basis
of a report by Roslyakov (1977), it is suspected that some
may survive (G. Archibald and S. Lantis 1979).
Oriental Crested Ibis populations in Korea are well docu-
mented, but the bird apparently never nested there (Gore
and Won 1971). L. Taczanowski (1891-93) reported 50
north of Seoul in 1888. There were no more reports until
1965 (Fisher et ah 1969:184-185). Individuals were dis-
covered there in the winter of 1977-78 by George Archibald
(Thiede 1982, G. Archibald and S. Lantis 1979). No con-
firmed reports exist after 1981 (W.W. Brehm-Fonds 1984).
In China, the populations apparently decreased gradu-
ally over the past century (Vaurie 1972). Oriental Crested
Ibises were unreported from China for over two decades,
from the late 1950s until 1972, when several were caught for
the Beijing Zoo. One captive ibis remained there as of 1984.
A government-sponsored survey in 1974 found no addi-
tional wild birds. After a 3-year search of eleven provinces
(20000km 2 ), a breeding group was finally located in the
Qinling (Tsingling) Mountains of central China in 1981 (Dr
Liu Yinzeng in Luthin 1984a).
As a result of the discoveries in Japan and China, in 1984
the known world population numbered 21 birds, including 1
captive in China and 3 in Japan. By 1989, it had grown to
49, including 3 captives in China.
ECOLOGY
The remaining ibises in China occur in forest patches in the
high valleys of the Qinling Mountains, 1200m above sea
level. The birds have been reported roosting in pairs
throughout the year. Such pairs choose trees near other
pairs for winter night roosting.
Oriental Crested Ibises feed by Probing in shallow open
water, especially in rice-paddies, freshwater ponds, lakes
and streams in forested areas. The presence of shallow water
may be crucial; in China, Dr Liu Yinzeng of the Institute of
Zoology in Beijing found that high water levels in streams
apparently decreased food availability during nesting
(W.W. Brehm-Fonds 1983). During the non-breeding sea-
son they feed in groups and frequently shift feeding sites
among rice paddies. Juveniles disperse in winter and forage
solitarily in lowland aquatic habitats. An upright posture is
assumed in alarm. In Japan the ibises appear to be rather
tame, especially the juveniles.
They feed on small fishes and molluscs. In China, they
feed on aquatic vertebrates and aquatic and terrestrial soil
invertebrates. The diet includes fish (carp, catfish, weather-
fish), frogs, newts, river crabs, crayfish, water-scavenger
beetles, diving beetles, beetle larvae, crickets, snails, mus-
sels and earthworms (Yinzeng in Luthin 1984a, Yamashina
1978). The diets change seasonally, and intake appears to
be reduced in winter.
BREEDING
Oriental Crested Ibises returned to their nesting area in
October in China and January in Japan. The remnant birds
in China nest only in forest patches in remote high moun-
tain valleys. These (and similarly high mountain birds in
Japan) nest solitarily. However, lowland populations in
central China and Japan apparently were colonial.
The birds appear to be monogamous, and, as noted
above, paired outside of the nesting season. Mutual preen-
ing is common within such pairs. It appears to be solicited
by an ibis nibbling the bill of its associate before lowering its
head to be preened (G. Archibald and S. Lantis 1979). A
rejection of the invitation to preen is indicated by a Head
Bobbing display.

During the breeding season, ibises preen in such a way as
to distribute black pigment over their feathers. As described
by Uchida (1970), daubing behaviour includes rubbing the
head and neck against the shoulder feathers for as many as
30 min per bout. This occurs after bathing, taking 5 or 6
daubing sessions to distribute the pigment.
It seems that the complete repertoire of nesting behaviour
is not yet known. Twig presentation with Head Bobbing
seems to be an important aspect of pair-formation and pair-
maintenance and may occur several months prior to nest-
ing. When no potential mate is receptive or present, a bird
may engage in Twig Carrying for many days. This behav-
iour often precedes a Mounting display in which the male
mounts the receptive female and shakes his tail without
actual copulation (G. Archibald and S. Lantis 1979). Such
pseudocopulation occurs frequently, many times per day
especially when other ibises approach the pair. Bill touching
and allopreening are common during courtship.
The nest, constructed of twigs, is 50-70 cm in diameter. It
is placed in a pine or oak tree, rather far out on a branch (G.
Archibald and S. Lantis 1979). The nest-cup is lined with
soft material, including leaves and mosses. The clutch is 2-4
brown-spotted blue-grey eggs, each weighing 65-75 g and
63-68 mm long. In 1983, one of the captive Chinese pairs
built several nests, but never laid any eggs.
Incubation begins in March in China and (formerly)
mid-April in Japan. Each pair defends its nesting territory.
Both sexes incubate for 28-30 days. When departing the
nest during incubation, the adults cover the eggs with the
nest-lining material. Hatching in China is in mid-May.
At hatching young are light grey, with a featherless head
and orange-red legs. In China, nestlings are fed by regurgi-
tation and fledge in June. Juveniles remain with the parents
near the nesting area until July in China and until early
September in Japan.
Known birds have had limited nesting success. In China,
Zing found in 1983 that crows destroyed clutches, but that
the ibises can renest. One of the captive birds in Japan died
from egg binding. Encouragingly, in 1989, the 6 pairs of
wild birds laid and hatched 7 eggs.
Survival of hatchlings depends on continued food avail-
ability (W.W. Brehm-Fohds 1983). The adults and young
appear to remain together for some time. Birds from the
Qinling Range move to the Hanjang River, 20km away
during the dry summer months, but the adults return with
their young to the breeding grounds in winter.
TAXONOMY
The interesting cosmetic coloration of this species caused
considerable confusion among early taxonomists. Pere
David obtained specimens of the grey form, naming them
Ibis sinensis. D.G. Elliot (1877), following Oustalet, believed
them to be young of the year. Yasuo Uchida definitively
explained this unique type of cosmetic coloration.
Perhaps owing to its coloration and the close relationships
between many plant and animal species of China and the
southeastern USA, the Oriental Crested Ibis is thought to
be most closely related to the American White Ibis (Eudoci-
Oriental Crested Ibis 251
mus). Of course, detailed morphological study is required to
address this supposition.
CONSERVATION
Both known recent nesting locations of the Oriental Crested
Ibis are in preserves set aside for its protection. The conser-
vation of this species is a matter of special concern, consider-
ing its precarious status. Individuals, conservation organiz-
ations, and the governments of Japan and China have been
actively involved in the efforts. The participants include the
Institute of Zoology in Beijing, the Toki Center in Japan,
the W.W. Brehm Fund for International Bird Conservation
at Vogelpark Walstrode, Germany, the Worldwide Fund for
Nature, the International Crane Foundation, and the
Storks, Ibises, and Spoonbills Specialists Group of the
I.C.B.P.
The Oriental Crested Ibis has for many years been a well-
known bird in Japan. The pale rose-coloured feathers were
used by Samurai for their ornamental arrows and for feather
brooms in the tea ceremony (Takashima 1957a). The de-
crease in this species was caused by hunting, loss of habitat
by deforestation, pesticides, and modern rice farming prac-
tices that remove water after crop harvest. After their dras-
tic reduction and rediscovery, they were made a Special
National Monument in 1952. The Toki Center was founded
in Japan to protect the Crested Ibis. A programme initiated
to attract ibises to uncontaminated sites in Japan failed to
influence the widely-ranging birds to avoid other feeding
sites. In 1966, a captive breeding programme was estab-
lished with 6 young ibises. One of these birds survived
through 1984. Additional birds were brought into captivity
in the early 1980s (G. Archibald 1981). More recently, eggs
laid in Japan appear to be infertile, accounting for the lack
of hatching there for over a decade. Because of poor success
of the captives, attention was turned for a time to providing
feeding sites in rice-paddies continuously stocked with fish
(G. Archibald and S. Lantis 1979) . . . to no avail.
In China, the nesting area is protected, and a research
station has been established nearby. Observational studies
were conducted and food provided to the birds. In 1989, the
first Oriental Crested Ibis was hatched in captivity in the
Bejing Zoo, and another was hatched in 1990 (Specialist
Group on Storks, Ibises and Spoonbills 1989, 1990).
In 1984, a Crested Ibis Workshop met in Beijing to
discuss a management plan for the species. It proposed a
three-point conservation plan for continued protection,
expanded research and captive propagation. This is being
undertaken at the Beijing Zoo in cooperation with the W.W.
Brehm Fund and Vogelpark Walsrode, Germany. The gov-
ernment of China issued commemorative postage stamps in
honour of the ibis, and funds generated from the sale of first-
day covers were to be used for the conservation effort (W.W.
Brehm-Fonds 1985). Despite these efforts, the conservation
situation with respect to this species could hardly be more
severe.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 321.
 Eurasian
Spoonbill
Platalea leucorodia Linnaeus

Subspecies:

Platalea leucorodia leucorodia Linnaeus

Platalea leucorodia Linnaeus, 1758, Syst. Nat., ed. 10, p. 139; based on 'The Spoonbill' of Albin,
1734, Nat. Hist. Birds, 2, p. 61, pi. 66: Europe: restricted to Sweden by Linnaeus, 1761, Fauna
Svecica, ed. 2, p. 57
Platalea leucorodia balsaci Naurois and Roux
Platalea leucorodia balsaci Naurois and Roux, 1974, Oiseau, 44, p. 77: Zira Island, Bane d'Arguin,
Mauritania

Platalea leucorodia archeri Neumann

Platalea leucorodia archeri Neumann, 1928, Journ. Ornith., 76, p. 783: Dahlak Island, Red Sea

Other names:

White Spoonbill, Black-legged Spoonbill, Common Spoonbill, Indian Spoonbill, Japanese Spoonbill
(English); Spatule blanche (French); Loffler (German); Lepelaar (Dutch); Espatula (Spanish);
Spatola (Italian); Skedstork (Swedish); Kolpitsa, Kolpik, Lopaten ['shovel'] (Russian); Abu
Malaqah ['father of the spoon'] (Arabic); Herasagi ['spatula heron'] (Japanese); Ta-pyi-lu, Ta-shao-
tsui (Chinese); Chamach baza, Chamcha (Hindi); Ghinta, Khunte bak (Bengali); Handi-alawa
(Sinhala)

IDENTIFICATION
The Eurasian Spoonbill is a medium-sized wading bird, standing about 60-85 cm tall, with a wing-span
of c. 115-145 cm. It is white, rather stout and has a long neck and legs and a black spatulate bill.
The adult's all-white plumage can become dingy with soiling. The black bill has irregular horn-grey
and black barring on the upper side and an individually variable amount of yellow near the tip. The legs
and feet are black. There is a narrow band of bare black skin at the base of the upper mandible and on
the lores to the eye. The naked chin and throat are mainly pale sulphur-yellow. The iris is dark red or
carmine.
The plumages of the sexes are the same, but they are dimorphic in size. Males have slightly larger
and heavier bodies, longer legs and longer bills, differences that are evident within wild flocks.
During the breeding season, the plumage is immaculately white and the crest becomes particularly
obvious, especially during displays. The crest consists of long narrow feathers (up to 12.0-15.Ocm in
males, 10.5-13.5 cm in females). A crescent-shaped gorget or collar, with a variable amount of yellowish
buff to cinnamon, develops on the lower foreneck; sometimes the crest is lightly tinged with the same
colour. The bare skin of the lower throat becomes a richer yellow, often with a reddish band bordering
the feathers (Cramp 1977:357).
The two African subspecies differ from the nominate race as follows. The smaller balsaci has a
completely black bill, or black with only traces of yellow; the yellow-buff on the chest is faint or absent
in breeding season. The subspecies archeri is similar to balsaci^ but smaller still.
254 Eurasian Spoonbill
At hatching, the bill, legs and feet are orange, becoming
pink-flesh after a few days. The bare skin around the eye is
blue; the naked chin, throat and base of upper mandible are
pink; the iris is grey. A nestling's first-down is sparse and
white with silky tips; the second-down is short and dense,
cream-white. Bills are not spatulate in young nestlings.
Rather they gradually thicken and become bulbous at the
tip and slightly decurved.
Juveniles are similar to non-breeding adults, except that
the bill is pinkish orange, becoming dark during the first
winter. Full bill-length is not attained until 3^6 months.
Legs are pinkish flesh coloured, later changing to slate. The
iris is grey-brown. Some outer primaries and primary
coverts have horn-black tips, and primaries, secondaries
and some other feathers have dark shafts, although this is
variable. Dark primary-tips and feather-shafts gradually
decrease as birds mature, though some individuals, in other-
wise full breeding plumage, retain them (Cramp 1977:357).
This is a generally silent bird, except during social dis-
plays at the nest, when a deep grunting or groaning 'Huh-
huh-huh' is given. These soft vocalizations can be heard
over short distances only. During Sparring and other fights,
various snapping sounds are produced by the bill's rapid
closure and by bills striking against each other. This is not
true bill-clattering, as occurs in most storks. Small nestlings
Beg with high-pitched 'Chittering' calls; in older nestlings
these calls become a lower-pitched 'Tschibb-tschibb-
tschirr-tschirr' (Bauer and Glutz von Blotzheim 1966:430).
Like all spoonbills, this species flies with neck and legs
extended. A series of wing-beats alternates with a short
glide. Flocks often fly evenly spaced in single file or in
diagonal wavy ribbons. They sometimes soar to consider-
able heights on thermals. The flapping rate has been
recorded as 186 ± 1.8/min (Palmer 1962:533).
The Eurasian Spoonbill is distinguished from the white
egrets by its novel bill, slightly heavier body and neck, and
characteristic feeding behaviour. In flight it can be separ-
ated from egrets by the extended neck and faster wing-beats.
In the southern parts of its range, it might be mistaken, at a
distance, for a white ibis, but the straight, spatulate bill of
the spoonbill is unmistakable. In Palearctic regions, it could
be mistaken, in flight at long range, for a swan (Cygnus), but
the spoonbill flaps and glides, and its long legs extend well
beyond the tail. In northern Africa, it is distinguished from

the African Spoonbill by the latter's bare red face, red legs,
red on the bill and pale blue iris.
DISTRIBUTION AND POPULATION
This spoonbill breeds from Europe to northeastern China,
India, Sri Lanka, and the eastern and western coasts of
northern Africa. It winters south to Mali, Sudan, Persian
Gulf, India, Sri Lanka, southeastern China, and southern
Japan.
Platalea leucorodia leucorodia formerly bred rather widely
throughout Europe. Until the 17th century it nested in
southeastern England. It now breeds more locally in Eur-
ope: southern Spain, Portugal, northwestern Netherlands,
Denmark (sporadically, north to 57°04'N; Kortegaard
1973), eastern Austria, Hungary, Greece, the Balkans and
southern USSR. Recently, tentative breeding attempts have
occurred again in France, northern Germany and Czechos-
lovakia. The range extends discontinuously across Asia:
from Turkey and southwestern Siberia, eastward to western
Mongolia, southeastern Siberia and northeastern China,
and southward to India and Sri Lanka. It is a regular visitor
to the UK, and casual or accidental in Scandinavia,
Madeira, Canary Islands, Cape Verde Islands, Faeroes,
Azores, and also Greenland (Cramp 1977:353).
Platalea leucorodia balsaci is resident on islands off the coast
of Mauritania, West Africa, primarily the Bane d'Arguin, at
about 20°N. It is probably non-migratory.
Platalea leucorodia archeri is resident on the coasts and
islands of the Red Sea, the Gulf of Aden and Socotra. It is
said to be a 'very common breeding resident' along the
northwestern coast of Somalia (Ash and Miskell 1983).
Several small colonies have recently been established north-
ward to Nabq, southern Sinai and Tiran Island at the
mouth of the Gulf of Aqaba (Paz 1987:36). This population
is probably non-migratory.
The wide-ranging leucorodia subspecies is migratory and
also undergoes marked disperals. European populations
winter mainly in Africa, regularly south to Mali and Sudan,
rarely to Nigeria, Kenya and Uganda. A few remain around
the Mediterranean bas,in, occasionally as far north as The
Netherlands and England in mild winters. Eastern Euro-
pean populations migrate via Greece to the Nile Valley in
Sudan and via Italy to Tunisia (Miiller 1984). Western
Asian populations winter from Israel, Iraq, Persian Gulf,
north coast of the Arabian Sea, and India southward to Sri
Lanka. Eastern Asian populations winter mainly in south-
eastern China, rarely to southern Japan and Taiwan.
It is mainly the temperate Palaearctic populations that
are migratory, whereas the more southerly populations are
primarily resident. The migration routes and timing are for
the most part well known. Migratory movements are
usually diurnal, in small parties or large flocks.
In the late summer there is a post-breeding dispersal,
mainly of juveniles, which generally occurs over short dis-
tances, although sometimes it is further, e.g. Yugoslavia to
Scotland, Hungary to the Camargue, France, and The
Netherlands to the Azores.
Birds depart European colonies on regular migration in
Eurasian Spoonbill 255
August-September and return in March-April. Colour-
ringed birds from The Netherlands are known to follow the
Atlantic coast of western Europe and have been found, in
winter, along the west coast of Africa in Senegal and Mauri-
tania. It is believed that many Dutch and Spanish birds
winter among the larger population of P.L balsaci in and just
south of the Bane d'Arguin, Mauritania (Roux 1962, 1973,
Poorter 1982); this has been confirmed by ringing recoveries
(Trotignon and Trotignon 1981).
Ringed birds from eastern Europe have been recovered in
winter in southeastern Europe, Italy, Turkey, Tunisia,
Libya, Egypt, Sudan and Niger. The virtual absence of
recoveries from the northwestern Mediterranean, eastern
Morocco and coastal Algeria suggest there is little contact
between eastern and western European birds on their win-
tering grounds. Birds ringed at the large colony at Lake
Manyas, Turkey, have been found in Israel, Egypt, Sudan,
Iraq and Pakistan (Schiiz 1957b). They more often utilize
marine habitats—such as deltas, estuaries, lagoons and
shallow inlets —as their winter quarters.
During the last century, vast flocks of migrant Eurasian
Spoonbills would arrive in India from the north during
October. More recently, 5 spoonbills, ringed as nestlings in
the Caspian and Black Sea regions of the USSR, were later
recovered in India (Sapetin 1968, McClure 1974:94).
Wetten (1986) estimated the world population of Eura-
sian Spoonbills to be 31 000-34500 individuals. The species
is abundant nowhere in Europe; in most countries where the
bird breeds there are only a few hundred pairs in the best of
years. Population trends in Europe have been generally
downward throughout this century, with only some very
recent cause for encouragement (G.A. Brouwer 1964, Bauer
and Glutz von Blotzheim 1966:432-435, Cramp 1977:353-
354). The history in The Netherlands is illustrative. From
300 pairs around 1900, the population rose with protection
to 450 pairs in 1925, and to about 500 pairs in the 1950s. It
decreased to below 200 pairs in 1964. In 1989, owing to
increased environmental protection, the population had
rebounded to 423 pairs nesting in 8 colonies (K. Brouwer
pers. comm. 1990). The first nesting of the Eurasian Spoon-
bill in France occurred in 1981 (Marion and Marion 1982).
In Spain during 1990, there were 850-900 nesting pairs in 2
colonies (E. Aguilera pers. comm. 1990). Tbe breeding
population in Austria and Hungary has remained almost
constant since 1950 (Miiller 1984). In contrast, it may have
disappeared completely as a breeding bird from Romania
(Wetten 1986).
The West African population (P.L balsaci) in the Bane
d'Arguin, Mauritania, may have the densest concentration
known for the species, with an estimated 1430 breeding
pairs during the years 1959-65 (de Naurois 1969:106) and a
total population of 5000-6000 birds in 1974 (Poorter 1982).
Between 8600 and 10000 spoonbills were counted in the
Bane d'Arguin area during December 1979 and January
1980 (Trotignon and Trotignon 1981); an unknown portion
of these were winter visitors from Europe.
The Red Sea population (P.L archeri) has the smallest
numbers, with an estimated 200-500 birds in eastern coastal
Sudan (Luthin 1984a).
256 Eurasian Spoonbill
During censuses in mid-winter 1989, 5559 Eurasian
Spoonbills were counted in India and Sri Lanka, the
greatest numbers being 1751 birds in Gujarat and 1065 in
Rajasthan (D.A. Scott and P.M. Rose 1989). It is not known
how many of these were migrants from further north and
how many were birds resident in the Indian region.
ECOLOGY
The habitat of the Eurasian Spoonbill is rather broad,
including many types of shallow-water areas with mud, clay
or fine-sand bottoms. Birds usually avoid waters with a
rocky bottom, thick vegetation or swift currents. They are
usually found in coastal lowlands or alluvial river valleys, in
fresh, brackish or salt water. They feed in estuarine areas,
especially on the wintering grounds. However, European
birds are relying increasingly on estuaries for feeding during
nesting (J. van Wetten in Luthin 1984a).
The birds feed both singly and gregariously in aggre-
gations of up to 100 birds, generally in water less than 30 cm
deep. Their foraging behaviour is characteristic of spoonbills.
Food detection is primarily non-visual. A bird wades vigor-
ously forward, Walking and Sweeping its partially open bill
from side to side in the water with a semi-circular scything
motion. The bill is gaped about 5 cm, and the tip often
touches the substrate. When active prey such as fish are being
hunted, birds may hold their bills to one side and run rapidly,
Dragging them through the water (Poorter 1969). When less
active prey is sought, birds usually walk more slowly and
search with wider sweeps of the bill (Wetten and Wintermans
1986:13). They also use an Intensive Sweeping, called terug-
zoeken (trying to relocate) by Poorter (1969), when they
remain searching in one place. Spoonbills exhibit a Probing
behaviour (wroeten—rooting; Poorter 1969), using a stabbing
motion. S. Ali and S.D. Ripley (1968:117) state that 'where
food is plentiful a compact, eager, jostling herd will advance
almost at a run, working methodically back and forth over a
particularly rewarding patch5.
Foraging is most active during mornings and evenings,
and like most spoonbills, this species often also feeds at
night (Aguilera 1990b). In coastal areas feeding is primarily
at low tide, regardless of the time of day (van Oordt in
Bannerman 1957:30).
The diet consists of small fish (up to 10-15 cm), aquatic
insects and their larvae, crustaceans, frogs and tadpoles,
molluscs, worms, leeches, reptiles, and some plant material
(possibly ingested accidentally). At freshwater feeding areas
in The Netherlands, spoonbills captured 1.5-2.0 prey items
(mostly small fish) per minute, i.e. 540-960 prey items
during a 6-8 h feeding day (Wetten and Wintermans
1986:44). The amount of food eaten has been recorded: 2
captive spoonbills together ate about 1 kg of freshwater fish
per day (O. Koenig 1952); an adult weighing 1.9kg ate
300 g/day, and a juvenile weighing 0.9 kg ate 140 g/day (I.
Sterbetz 1984). Although the usual prey is small, there is a
report of a bird in The Netherlands that choked while
attempting to swallow a large eel (R.P. Allen 1942:101).
During the breeding season, feeding areas are usually
within 35-40 km of the nesting colony (Wetten and Winter-
mans 1986:47). They often walk ashore after feeding and
rest on one leg. Spoonbills commonly sleep with their bill
and face rotated to the rear and tucked beneath the feathers
of the back. They roost either on the ground (if secure) or in
trees. They can swim well, and do so when necessary.
BREEDING
The Eurasian Spoonbill usually nests in colonies of a few to
a few hundred pairs. They nest in dense stands of emergent
aquatic vegetation, especially reeds and trees, such as wil-
lows (Salix sp.), oaks (Quercus sp.) and poplars (Populus sp.)
(Beetham 1910, Marion and Marion 1982, Miiller 1983,
Aguilera and Alvarez 1989). Breeding colonies often face
pools or channels. Almost invariably the nest site is within
10-15 km (often much less) of feeding areas. Nests are
usually at low elevations, but an exceptional colony was at
nearly 2000 meters on Lake Sevan, Armenia (Dementiev
and Gladkov 1951:417, Cramp 1977:352).
Sometimes Eurasian Spoonbills nest in or near mixed
colonies of herons, ibises and other waterbirds, but they
usually remain somewhat segregated from other species. A
colony site is often used in successive years. In most areas
birds are intolerant of disturbance, especially early in the
breeding cycle; however, they often nest in trees in the midst
of villages in India (E.C.S. Baker 1935:433).
Nests are usually placed 1-2 m apart, but sometimes so
closely together that they touch. They are built in trees (up
to 2-5 m, sometimes higher), reedbeds or on the ground, as
is the case in The Netherlands and Mauritania. The nest
itself is a large pile of reeds, sticks or twigs, lined with grass
and leaves, 50-100 cm in diameter and 30-60 cm deep.
Pair-formation displays, described by Poorter (in Cramp
1977:355-356), appear to be similar to many of those seen in
the African and Royal Spoonbills (Kahl 1983, 1988a).
During the Greeting display, the brightly coloured throat-
skin is exposed to the mate when the head is raised. The
luxuriant crest is raised and fanned laterally into a 'war-
bonnet5 during many behaviours. Sparring, Nest-Covering
displays and Begging displays also occur. Mutual allopreen-
ing by mates is common, especially on the head and neck,
during quiet moments on the nest.
Copulations are frequent in the early breeding season: as
many as 11 attempts in 1 h have been observed (Dragesco
1961b). The average interval between the first observed
copulation and the day on which the first egg is laid is
6.7 days (n = 9) (Aguilera and Alvarez 1989). Attempted
extrapair copulations by paired males with neighbouring
females, while their mates are absent, are common (Agui-
lera 1989). During copulation, we observed that the male
rapidly bites and slaps the bill of the female, making a soft
clattering or slapping sound (pers. obs. M.P.K.). In 18
copulations timed by Aguilera and Alvarez (1989) the dur-
ation was 9.7 ± 2.3 s. The same authors mentioned, but did
not describe, a pre-copulatory display by the male, termed
'Rubbing back5.
Each pair defends its own nest and the immediate area
nearby, in which only close neighbours are tolerated. Birds
from adjacent nests often cooperate to drive away strange

adults. Both sexes assist in the construction of the nest, with
males collecting most of the nest material at Spanish nests
(Aguilera and Alvarez 1989).
Eggs are slightly elongated ovals, chalky white with small
reddish brown spots and lines. It was found in Romania
that larger clutches have smaller eggs (Vespremeanu
1968a). The clutch size is usually 3-4, rarely 6 and excep-
tionally 7 (de Naurois 1959). In Macedonia, 54 clutches
were 3 eggs (56% of clutches), 4 (37%), 5 (6%), and 6
(1%). In Romania, 415 clutches were 2 eggs (23%), 3
(54%), and 4 (23%), with a mean of 3.0 (Vespremeanu
1968a). In southwestern Spain a fifth egg was laid in only
3% of 937 clutches (Aguilera 1990a).
Eggs are laid at 2-3 day intervals (Cramp 1977:356). The
incubation period is given by Holstein (1929) as 21 days,
but it is probably nearer to 24—25 days (Dementiev and
Gladkov 1951:419, Vespremeanu 1968a). Incubation is by
both sexes and usually begins with the laying of the second
egg; hatching is asynchronous (Aguilera 1990a). Pairs
within a colony may be relatively unsynchronized, as all
stages of nesting may occur simultaneously (Vespremeanu
1968a). At a colony in southwestern Spain, males generally
attended the nests in the daytime and foraged at night,
whereas females tended the nests mostly at night (Aguilera
1990b, Aguilera and Alvarez 1989). Eggs and small nes-
tlings are often shaded by their parents on hot days.
At feeding a nestling inserts its bill into the buccal cavity
and upper throat of the adult to receive food that is regurgi-
tated for it. Unfledged young wander about the colony from
about 3 weeks of age, often visiting other occupied nests;
they return to their own nest for feeding up until about 6-8
weeks. The fledging period is 45-54 days (Vespremeanu
1968a, Cramp 1977:356). Both parents tend the young until
well after fledging, females were observed to feed the chicks
more than males at Spanish nests (Aguilera and Alvarez,
1989). The bond between parents and young may continue
until about 10 weeks, with the parents gradually providing
fewer meals and the young foraging more on their own.
Most nestling mortality occurs in the first 10-15 days.
Breeding success was recorded by Vespremeanu (1968a) in
Romania in 1958-63: of 1244 eggs laid, 95% hatched, and of
1188 young, 70.3% fledged. Nestling mortality varied from
11.3 to 100.0%, depending on the season. At the Goto Donana
National Park, Spain, 4 is the maximum known number of
young fledged from any spoonbill nest (Aguilera 1990a).
In broods of 5, the death of the youngest chick is some-
times attributable to attacks by their own parents, rather
than by siblings (Aguilera 1990a). High water levels during
the nesting season can lead to very low success (Vespre-
meanu 1968a). At some European colonies, food availability
may be a limiting factor, because adults have to fly longer
distances (often as much as 30 km) to feed as the season
progresses (J. van Wetten in Luthin 1984a).
Avian predators, such as Jackdaws (Corvus monedula),
Magpies (Pica pica), and Black Kites (Milvus migrant), prey
on eggs and young in Spain (Aguilera and Alvarez 1989). At
other sites mammalian predators account for much of the
egg and nestling mortality, except on offshore islands that
are out of reach of terrestrial predators.
Eurasian Spoonbill 257
TAXONOMY
Some authors have considered the Royal Spoonbill, Platalea
regia to be a subspecies of the Eurasian Spoonbill (i.e.
Platalea leucorodia regia). However, based on field studies of
the various populations on their breeding grounds (Kahl
1988a and pers. obs.), we conclude that the two are distinct
enough to warrant specific status.
There have been suggestions that the eastern and western
populations could be considered subspecifically distinct. We
agree with earlier statements (Seebohm 1890:230, Meinertz-
hagen 1954:394, Vaurie 1965:78) that the size differences
used to separate the eastern and western populations in
Eurasia are not sufficient to warrant the recognition of the
far-eastern race of P.I. major. We do, however, recognize
three other subspecies, based on differences in size and
breeding coloration: P. 1. leucorodia, P.I. balsaci and P.I. archeri.
It seems that P. leucorodia, P. regia and P. minor are suf-
ficiently closely related to comprise a superspecies. But P.
alba is more distinctly different from the other three and not
so closely similar as indicated by some authors (e.g. Snow
1978:42).
CONSERVATION
The breeding range of the Eurasian Spoonbill, especially in
western Europe, has been drastically reduced owing to
drainage, alteration or pollution of feeding areas and,
mainly in earlier years, to human exploitation of eggs and
nestlings for food (G.A. Brouwer 1964). An important con-
sideration over the past century has been the disappearance
of reed swamps, the favourite nesting habitat, due to agri-
culture and hydropower development (Vespremeanu
1968a).
The decrease of the Dutch population from about 500
pairs in the 1950s to 150-200 pairs in the early 1970s,
probably owes to pollution of feeding areas by chlorinated
hydrocarbons (Rooth and Jonkers 1972). Dutch birds re-
covered to about 400 pairs in the late 1980s, and by 1989
there were 423 pairs in 8 Dutch colonies (Osieck and
deVries 1987; K. Brouwer pers. comm. 1989). Populations
in Greece continue to decrease owing to degradation of
habitat, agricultural practices, overfishing and disturbance
(Crivelli et al. 1988). The species is now seriously endan-
gered in most areas of its European range (Cramp
1977:353).
During their migrations, the western European popu-
lations are dependent on a succession of estuaries along the
coasts of Belgium, France, Portugal, Spain and Morocco.
Eastern European birds depend on sites in Italy and Greece.
It is important that these migratory and staging areas are
protected, as well as feeding and nesting habitats in the
summer and winter ranges.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
pages 322-323.
/v
 Royal Spoonbill
Platalea regia Gould

Platalea regia Gould, 1837, Proc. Zool. Soc., pt. v, p. 106: east coast of New South Wales

Other names:

Blackbilled Spoonbill, Blackfaced Spoonbill (English)

IDENTIFICATION
The Royal Spoonbill is a medium-sized, rather stout, white waterbird, standing 74-85 cm tall, with a
wing-span of about 120 cm. It has a long neck and legs and a black, flattened bill, spatulate at the tip.
Although the plumage of the adult is entirely white, it is sometimes soiled. The bill is mostly black,
with irregular bluish grey and black barring on the upper side; it lacks the yellow tip seen in Eurasian
Spoonbills. The skin of the face and unfeathered throat is black. The legs and feet are shiny black. The
iris is a deep, dark crimson.
The species is sexually dimorphic. Males have slightly larger bodies, longer legs, and longer bills
(Vestjens 1975b, Kahl 1988a).
During breeding the plumage is immaculately white, and the nuchal crest becomes prominent. It is
more luxuriant than in any other species of spoonbill, composed of up to 100 long, narrow plumes
(individual feathers measured up to 19.4 cm long). The crest is especially conspicuous when erected and
fanned laterally during displays. Some adults begin losing their crest feathers even before the young
have left the nest. Also during breeding, a horizontal collar of pale yellowish buff appears on the lower
foreneck; this is variable and less conspicuous than in the Eurasian Spoonbill. Some individuals have a
faint hint of the same buff colour on their crest. A naked, narrow band of bare skin under the wings is
salmon-pink to saffron coloured (Kahl 1988a).
Oval spots, about 20 X 10 mm, of deep yellow-ochre develop on the skin above (and sometimes, to a
lesser extent, below) the eyes, contrasting strongly with the black facial skin. Also a triangular patch of
dull red skin appears at the top of the black forehead, just below the white feathers of the crown. These
yellow and red facial markings are variable between individuals. According to K.W. Lowe (pers. comm.
1990) some Royal Spoonbills can be seen at all months of the year with these breeding colours. (In
preserved specimens, the yellow eye-spots and red forehead-triangles of living Royal Spoonbills both
appear as dull yellowish tan areas. This has resulted in misleading descriptions by those unfamiliar with
the living birds.)
The bill, legs and feet are pinkish orange at hatching; the bare skin around the eye is bluish. The first-
down is sparse and white; the second-down is more dense and off-white. Bills are not spatulate in
nestlings but gradually thicken and become bulbous at the tip, with a slight decurvation (Kahl 1987c).
Juveniles are similar to non-breeding adults, except that the bill is shorter and dusky pinkish, becoming
longer and darker during the first year. The facial skin is dusky and lacks any coloured areas, becoming
blackish during the first year. The legs change gradually to slate-grey. In certain individuals, some
outer primaries and primary coverts have horn-black tips, and primaries, secondaries and some other
feathers have dark shafts. Dark primary-tips and feather-shafts gradually fade or disappear as the birds
mature. However, some apparent adults retain dusky primary-tips.
The Royal Spoonbill is generally silent, except during social displays at the nest, when a low
growling, grunting, or honking *Huh-huh-huh-hutf is given (Kahl 1988a). These vocalizations are soft and
audible only over short distances. During fights, some rattling sounds are produced by snapping of the
bill. True clattering, as made by storks, is not heard, contrary to some reports in the literature. Small
260 Royal Spoonbill
nestlings perform the Begging display with high-pitched
'Chittering* calls; in older nestlings these calls become
lower-pitched (Kahl 1988a).Vestjens (1975a) found that the
shape of the trachea changes with age, and this may be
related to changes in voice.
Flight is with the neck and legs extended, a series of wing-
beats alternating with a short glide. Flocks often fly in single
file or in diagonal wavy ribbons, with each bird evenly
spaced behind and to one side of the one in front. The
flapping rate averages 230/min (n = 44) (pers. obs.
M.P.K.).
The Royal Spoonbill is distinguished from the white
egrets by a longer, heavier and spatulate bill, slightly hea-
vier body and thicker neck, characteristic feeding behav-
iour, and, in flight, by an extended neck and faster wing-
beats. It might be mistaken at a distance for an Australian
White Ibis, but the straight, spatulate bill is diagnostic. It is
distinguished from the slightly larger Yeilowbilled Spoonbill
by the black face, bill and legs. The Royal Spoonbill usually
has a whiter, 'cleaner' plumage than either the Australian
White Ibis or the Yeilowbilled Spoonbill. In the extreme
northwestern part of the Royal Spoonbill's range (e.g. Bor-
neo), it may rarely overlap with wintering Blackfaced
Spoonbills. In non-breeding plumage, it might not be poss-
ible to distinguish this bird from the Blackfaced Spoonbill,
as discussed in that species' account.
DISTRIBUTION AND POPULATION
The Royal Spoonbill is primarily Australian, found in the
well-watered parts of northern, eastern and southeastern
Australia. In Indonesia, it is possibly resident in the Mollu-
cas, Lesser Sunda Islands, Sulawesi and Irian Jaya; it has
occurred in Java, where it formerly bred (Hoogerwerf 1952,
Silvius and Verheugt 1990). In southern Papua New
Guinea, it is commonest in the dry season and probably
only a non-breeding visitor from Australia; a small, isolated
population exists on Rennell Island, Solomon Islands
(Coates 1985:90, Beehler et al. 1986:61). Prior to 1950, it
was a vagrant in New Zealand, but more recently it has
bred in small numbers at Okarito and Otago, South Island
(Holdaway 1980, H.A. Robertson and B.E. Preece 1980). It
wanders to Tasmania (Blakers et al. 1984:66) and Borneo
(Smythies 1981:33).
This species is sedentary or nomadic, moving mainly in
response to periodic droughts. However, it is reported to
migrate across the Torres Straits, between Australia and
New Guinea, in large flocks (Coates 1985:90). Five recover-
ies, 1-36 months after ringing, show movements of between
277 and 1472km within Australia (Blakers et al. 1984:66,
K.W. Lowe, Australian Bird and Bat Banding Schemes
pers. comm.).
Few estimates of population numbers and trends are
available. However, this is a common to locally abundant
species in the well-watered parts of northern and southeast-
ern Australia. In 1988, a dry-season survey by the Asian
Wetland Bureau found over 250 birds in southeastern Irian
Jaya (Silvius and Verheugt 1990). Except for these New
Guinea birds, possibly, populations outside Australia are
not large.
ECOLOGY
The Royal Spoonbill inhabits large shallow waters, inland
and coastal marshes, lake shores, mud flats and mangroves.
It does frequent farm dams, but is less often found on such
small waters than is the Yeilowbilled Spoonbill. In tidal
areas most feeding occurs at low tide.
Royal Spoonbills are often found in small flocks. They
habitually feed in water less than 40 cm deep. They rest on
the shore or in trees, with bills rotated about 150° and
tucked into the feathers of the upper back.
Feeding occurs both day and night. As do other spoon-
bills, the Royal Spoonbill feeds by touch. The partly open
spatulate bill is swept from side to side in the water.
Although the birds feed mostly by Slow Sweeping, they also
do a rapid, short-stroked Intensive Search, after detecting a
potential prey item. Vestjens (I975b), who described the
feeding behaviour in detail, found that this species also used
Dragging, Probing and Grabbing foraging methods. All
these techniques involve tactolocation of food, except for
Grabbing, in which prey is visually located. It is apparently
the food items that touch the widest portion of the 'spoon'
that are detected and grasped. Food is then transferred from
the distal portion of the bill to the throat with a quick,
backward toss of the head and then swallowed.
They are specialized and rather efficient foragers. During
'Slow Sweeping' foraging birds captured an average of 6.4
items per minute in the freshwater Lake Cowal, NSW
(Vestjens 1975b), and 3.5/min in the marine tidal area of
Westernport Bay (K.W. Lowe 1982). Spoonbill predation of
intertidal shrimps has been shown by R.K. Howard and
K.W. Lowe (1984) to be sufficient to influence their popu-
lation levels. The results of this study demonstrate how a
non-visual predator can be selective in the prey it catches.
Spoonbills were particularly selective for adult female
shrimps rather than small shrimps.
The diet includes small fish, aquatic insects, crustaceans,
small freshwater snails, tadpoles, and, occasionally, spiders
and crickets (Vestjens 1975b). Prey caught at night is differ-
ent to that caught during the day (K.W. Lowe 1982).

When feeding in the same habitat, the Royal Spoonbill
takes more fish and fewer crustaceans and insects than does
the Yellowbilled Spoonbill; this is due to differences be-
tween the two species in bill-structure and speed of move-
ment. In the stomachs of 20 Royal Spoonbills at Lake
Cowal, NSW, Australia, the ratio of crustaceans and insects
to fish was 4:3 (Vestjens, 1975b). At Westernport Bay,
Victoria, Australia, the stomach contents of 10 birds was
73% crustaceans (wet-weight), especially the striped prawn
(Macrobrachium intermedium), and 23% fish, especially (K.W.
Lowe 1982) the Bridled Goby (Arenigobius bifrenatus).
BREEDING
The Royal Spoonbill nests singly or in small colonies (up to
100 pairs), often with ibises, darters or the smaller cormor-
ants. The nest-site is usually within 5 km of feeding areas,
sometimes as near as 500 m (Vestjens 1975b).
The nest is a shallow platform of sticks in trees or bushes;
it is usually over water, occasionally up to 20m high,
although usually considerably lower. At Lake Cowal, nests
were placed in medium-sized trees of river cooba (Acacia
stenophylla), river red gum (Eucalyptus camaldulensis) or emerg-
ent lignum bushes (Muehlenbeckia cunninghamii). Nests are
sometimes constructed of reeds in reed-beds (Beruldsen
1980:158). In the small colonies at the periphery of the
range in New Zealand, nests were built in the tops of very
tall trees (at Okarito) or on the ground (at Blenheim)
(Holdaway 1980).
The timing of nesting is somewhat variable: August-
January in southern Australia and February-May in north-
ern Australia. On South Island, New Zealand, nesting is in
November-December, and in western Java was formerly
March-September.
Unmated birds of both sexes congregate in 'bachelor
parties', where many forms of ritualized displays are seen.
Males then establish themselves on potential nest-sites, and
females attempt to approach them. However, the females
are repeatedly driven away, with Supplanting or overt
attacks. Eventually a female is accepted by a male, and the
pair proceeds to build a nest. Mutual allopreening is com-
mon between mated birds, especially of the feathers of the
head and upper neck. Whenever one member of the pair
returns to the nest after an absence, both birds perform the
Greeting to each other. Other commonly performed ritua-
lized displays during courtship are Bowing displays (shown
by females), Sparring, Head Shaking, Stick Shaking, Dis-
play Sleeping, Display Shake, and Display Preen (shown by
both sexes). Copulations and egg-laying follow closely after
pair-formation (Kahl 1988a).
Both adults build the nest, with the male gathering most
of the material. The clutch size varies from 2-5 eggs, usually
3-A. The eggs are tapered oval in shape; they are dull white,
spotted with yellow-brown or red-brown. Incubation is per-
formed by both sexes for about 25 days; hatching is asynch-
ronous at 1- to 3-day intervals. Parents may desert the nest
if disturbed during the early stages of the breeding cycle,
especially during nest-construction before the eggs are laid
(pers. obs. M.P.K.).
Royal Spoonbill 261
Both parents brood, guard and feed the nestlings. At
feeding a nestling inserts its bill into the throat of its parent,
who then regurgitates food from the stomach for the young
to eat. Only one young is able to feed from the parent's
throat at a time. During the 1984-85 season at Lake Cowal,
a total of 110 nestling feedings averaged 6.9s in duration
(range = 4.7-12.5s); a brood was given between 8 and 28
individual portions during a session, and most of the feed-
ings took place within 10-20 min of the parent returning
from a foraging trip (pers. obs. M.P.K.). Young are capable
of their first flights at 4-5 weeks, but they continue to return
to the nest for feedings by the parents until about 2 months.
Family groups remain together for several weeks more.
Young have been observed performing the Begging display
on the wing, while flying after an adult (Hoogerwerf 1952).
Egg and nestling mortality are most commonly caused by
inclement weather or predation by Australian Ravens (Cor-
vus coronoides), lace monitors (Varanus varius) and, possibly,
various snakes (Vestjens 1977a).
TAXONOMY
According to some earlier classifications (e.g. J. Steinbacher
1979), the Royal Spoonbill was considered a subspecies of
the Eurasian Spoonbill, Platalea leucorodia regia. However,
owing to significant morphological differences, especially in
soft-part coloration, we prefer to treat it as a full species,
Platalea regia (Amadon and Woolfenden 1952, Gyldenstolpe
1955a,b).
If the Royal Spoonbill were conspecific with any other
population, it would be with the Blackfaced Spoonbill (P.
minor) of northeastern Asia, with which it shares many
morphological traits. As can be seen in the tables of body
measurements (see Appendix), there is considerable overlap
in size between the two populations, and the soft-part color-
ation is strikingly similar (see Ogilvie-Grant, 1889:48, col.
pi. 1). The extent and shape of the bare skin on the throat,
which has been suggested as a means to distinguish between
the Royal and Blackfaced Spoonbills, is quite variable,
individually and with age. It is, therefore, unreliable as a
taxonomic character. Pending further study, we chose to
retain Platalea minor as a full species; however, it is certainly
very close, taxonomically, to Platalea regia.
In our opinion, the three spoonbills, Platalea regia, P. minor
and P. leucorodia, are closely enough related to comprise a
single superspecies.
CONSERVATION
There appear to be only a few Royal Spoonbill breeding
colonies of significant size. However, in general, the species
seems to be doing well in Australia, the stronghold of its
range, and expanding slowly in outlying areas such as New
Zealand.
Status in the northern part of the range is less well known.
The species has apparently disappeared from Java, where it
nested in small numbers in the 1950s (Hoogerwerf 1952).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 324.
 Blackfaced
Spoonbill
Platalea minor Temminck and Schlegel

Platalea minor Temminck and Schlegel, 1849, in Siebold's Fauna Japonica, Aves, p. 120, pi. 76: Japan

Other names:
Lesser Spoonbill; Swinhoe's Blackfaced Spoonbill (English); Petite Spatule du Japon (French);
Kurotsura-herasagi (Japanese); Hsiao-pyi-lu, Hsiao-shao-tsui (Chinese)

IDENTIFICATION
The Blackfaced Spoonbill is a medium-sized white waterbird, c. 60-83 cm tall. As the common name
suggests, it has a black face, and the long bill is flattened and ends in a broad spoon.
The adult plumage is entirely white. The bare black skin on the face extends to the throat and 5-15
mm behind the eye. The bare area in back of the eye and on the throat is quite variable, and it is difficult
to judge in the field (pers. obs. M.P.K.). The bill is slate-coloured and, as described by Sharpe
(1898:51), 'transversely barred with black, the bars broken and disconnected on the spatule'. The legs
are black. The iris is crimson to blood red.
Sharpe (1898:51) also reported 'a patch and scattered spots of orange-ochre over the spatule of the
bill. Peter Kennerley (pers. comm. 1990) stated that this coloration on the bill was 'never seen on adults
in Hong Kong, even in nuptial plumage'. We wonder if Sharpe may have confused this species with the
Eurasian Spoonbill. If the Blackfaced Spoonbill does, indeed, show orange or yellow near the end of the
bill, it apparently does not occur in many individuals, nor to as great an extent as in the Eurasian. C.
Poole (pers. comm. 1990) tells us that birds 'can show pale areas on the spatule, which I would best
describe as fleshed coloured'; this description may refer to subadult birds. The matter of bill coloration
is one that requires additional observation.
Females are similar to males but slightly smaller. The plumage and soft-part colours change during
breeding, but the details remain somewhat uncertain. A yellowish half-moon marking develops on the
face, but its size, shape and position can vary. H.G. Won (1966) described the marking as being on the
lores. Referring to the facial spot, others place the 'bright yellow-ochre patch before the eye, extending
[across] the under-lid, and a thin line [across] the upper lid' (Sharpe 1898:51) or 'a yellow spot in front
of and under the eye' (La Touche 1931-34:432). The yellow eye-spots of living spoonbills appear as dull
yellowish tan in preserved specimens.
Breeding adults develop a long nuchal crest. According to Peter Kennerley (pers. comm. 1989) the
crest plumes of birds in Hong Kong take on a distinctive golden-yellow colour in the spring, just before
they begin migration. They also show a bright yellow bar across the lower foreneck or upper breast (not
on the throat, as described by H.G. Won 1966). In some cases, this develops into a complete golden
collar around the lower neck. In full breeding plumage the yellow collar of the Blackfaced Spoonbill is a
richer colour than the similar collar of the Eurasian Spoonbill (P. Kennerley pers. comm. 1990). Colour
photos in Chung (1986) show that by late in the breeding season, after the young have hatched, the
yellow marking on the lower foreneck has become faint, and the crest shows only a wash of pale buff.
The hatchling is little known, except that 'chicks bills are shorter than adults' and are round' (Chung
264 Blackfaced Spoonbill
1986) [from a translation by Kyoko Archibald]. The juven-
ile is white; it 'differs from the adults in having a more
yellowish bill and blackish shaft-stripes to the quills, the
outer primaries blackish at the ends of the outer webs'
(Sharpe 1898:51). These dark primary tips are obvious in
the field, making it easy to separate juveniles from adults in
flying groups.
Vocalizations of the Blackfaced Spoonbill have not been
described.
Birds we observed briefly in Hong Kong appeared to have
a flight typical of other spoonbills. The flapping rate which
was timed on a single occasion was 231/min (pers. obs.
M.P.K.).
This species is similar to the Eurasian Spoonbill but
noticeably smaller, with more extensive bare black skin
on the face and throat, and usually, perhaps always, lack-
ing the prominent yellow mark on the 'spoon' of the bill.
In its summer plumage this species shows more yellow or
buff on the crest than does the Eurasian Spoonbill (Gore
and Won 1971:123). It is possible that the Blackfaced and
Royal Spoonbills may occur together in Borneo (Mann
1989). For the reasons discussed below (see Taxonomy),
separation in the field would be difficult. In breeding plu-
mage (unlikely in Borneo) the Blackfaced Spoonbill would
have more yellow below the eyes, whereas the Royal Spoon-
bill would have more yellow above the eyes. The Blackfaced
Spoonbill may also lack the dull red triangle at the top of the
bill.
DISTRIBUTION AND POPULATION
The breeding distribution of the Blackfaced Spoonbill is
poorly known. It is confirmed to nest only in Korea, on a
few small islands (Sogam-do, Tegam-do, Tok-to; formerly
also on Ai, Kamsang) along the western coast of North
Korea (Chung 1986, Sonobe and Izawa 1987:36-37). Prior
to the Korean War (1950) this species was said to be
common and was found breeding on islands (e.g. Wido, Ito)
along the western coast of South Korea (Austin 1948:49, P.
Won 1973:26). There are unconfirmed reports of breeding
in central Manchuria in the vicinity of Harbin (Vaurie
1965:78), northwestern Jilin Province, Inner Mongolia,
and, possibly (formerly at least), in eastern China south to
Fukien.
As far as is known, most of the population winters in
southern China (Kwangtung Province, Hainan Island),
Hong Kong (Mai Po marshes), northern Vietnam (Hong
River delta) and southwestern Taiwan (Tsen-wen Hsi,
Chiayi Hsien). It has been recorded as far north as Korea in
December (H.G. Won 1966, Gore and Won 1971:123). [The
Eurasian Spoonbill was said 'to be numerous in the man-
groves' of coastal Vietnam in winter (Delacour and
Jabouille 1925), but this may have referred to this species
instead.]
A few winter stragglers are found in Thailand (P. Round
pers. comm. 1989), Japan (Takara 1979, Sonobe and Izawa
1987:36), and the Philippines (duPont 1971:23). A record
from Brunei was assigned to this species, but it was appar-
ently a subadult bird and could conceivably have been a
Royal Spoonbill from the south (Mann 1989, pers. comm.
1990). There may be other, as yet undiscovered, small flocks
wintering along the south China coast; probably few winter
inland in China (D.A. Scott pers. comm. 1990).
Wintering birds are present in Hong Kong mainly be-
tween late October and early May (Chalmers 1986:45); they
are found in Taiwan from September through May (Lucia
Liu Severinghaus pers. comm. 1990). According to H.G.
Won (1966) summering birds arrive in North Korea in
March (although this must be exceptionally early) and
leave again in October/November.
Migrating birds must pass through China, but little
is known about their movements between the breeding
areas in North Korea and the wintering grounds. Some do
cross South Korea, where up to 46 were recorded on South
Kanghwa Island on 5 September 1989; 20 were still present
into the second week of October 1989. About two-thirds
of these birds were immatures (C. Poole pers. comm.
1990). An undetermined number of Blackfaced Spoonbills
were identified in flocks of Eurasian Spoonbills during
December 1985 and January 1986 at Poyang Lake, Jiangxi
Province, China (Kennerley 1987), and single birds were
seen there also in December 1989 (P. Kennerley pers.
comm. 1990).
The known population of the Blackfaced Spoonbill is
small, and it is currently endangered. Information on the
breeding population is scarce. In June 1965, a total of 36
adults were counted at a breeding colony off the coast of
North Korea (Choi 1966); in the mid-1980s about 30 were

counted in that area (Chung 1986, Sonobe and Izawa
1987:37).
More information is available from the wintering
grounds. Prior to 1982 the highest winter counts in Hong
Kong were about 20 birds; however, since then, there has
been a slow but steady increase (Chalmers, 1986:45). In
most recent years about 40 birds have wintered in Hong
Kong. A high of 47 was recorded on 28 January 1989 (P.
Kennerley pers. comm. 1990). Of the 40 Blackfaced Spoon-
bills we observed in Hong Kong on 31 January 1989,
approximately 30-40% were immature birds (pers. obs.
M.P.K.).
Between September 1988 and March 1989 a group of 130
Blackfaced Spoonbills wintered in southwestern Taiwan
(D.A. Scott and P.M. Rose 1989:80, Lucia Liu Sever-
inghaus pers. comm. 1990). Approximately 145 wintered
there in 1989-90 (Lucia Liu Severinghaus pers. comm.
1990), and about 100 were counted in the area on 12 March
1990 (C. Poole pers. comm. 1990).
In the winter of 1987-88 a total of 62 were counted, by Dr
Le Dien Due, in the Hong River delta area of Vietnam. A
high count of 27 birds was recorded there on 27-28 March
1990. Over 50% of birds seen in Vietnam during 1988 and
1989 were immatures (D.A. Scott pers. comm. 1990).
A few Blackfaced Spoonbills have recently been reported
during the winter in South Korea (Coulter and Brouwer
1991). 3-6 birds have wintered at Songsanp'o on Cheju
Island, and 4 were reported in 1987 in the Nakdong Estu-
ary.
If the Hong Kong, Taiwan and Vietnam populations
represent most of the wintering Blackfaced Spoonbills, the
total world population is, perhaps, about 285 birds (Ken-
nerley 1990). However, only c. 30 adults are known to
summer in North Korea. So it remains to be discovered
where the others go. Perhaps there are as yet undiscovered
breeding areas in the eastern part of China. The most likely
area would seem to be the offshore islands in Korea Bay, to
the east of Dalian, China, and adjacent to the North Korean
border.
ECOLOGY
The ecology of the Blackfaced Spoonbill is little known. Like
other spoonbills, it frequents shallow waters (ptchecopar
and Hue 1978:80). The known population occurs largely in
coastal areas, inhabiting tidal flats, saltmarshes, estuaries
and tidal creeks; it sometimes uses inland lakes. It appears
to favour mangrove areas during the winter.
The feeding behaviour is described as similar to other
spoonbills. The bird places the partly open bill in water,
Swinging it from side to side (Etchecopar and Hue 1978:80,
Chung 1986). According to H.G. Won (1966), 'In the sea-
sons when they were not breeding, they search [for] food in
the shallow waters along the seashore at dawn or late
evening (frequently in creeks and streams of tidewater)'. It
is 'met with in small parties, fishing in the shallows' (La
Touche 1931-34:433). Foraging birds seen on the wintering
grounds in Vietnam were mainly solitary (D.A. Scott pers.
comm. 1990).
Blackfaced Spoonbill 265
Blackfaced Spoonbills eat small fish, shrimps, tiny crabs,
snails and insect larvae (H.G. Won 1966). An immature
female collected in southern South Korea in December 1959
had 'many small fresh-water shrimp' in its stomach (Fennell
and King 1964). Delacour and Jabouille (1931, cited in R.P.
Allen 1942:101) reported that on the wintering grounds, in
what then was French Indo-China, they fed on insects,
crustaceans and vegetation.
The general behaviour (comfort movements, resting,
flight) that we observed in wintering birds in Hong Kong
did not differ noticeably from that of the other spoonbills
(pers. obs. M.P.K.). On the wintering grounds in Vietnam,
birds gather in groups only at high-tide to roost (D.A. Scott
pers. comm. 1990). They 'often stand on one foot with their
necks on the backs when roosting' (H.G. Won 1966).
BREEDING
The nesting biology of this spoonbill is poorly known. Nest-
ing is in June and July in Korea. All the presently known
breeding sites are on small, rocky offshore islands. From the
descriptions and photographs in H.G. Won (1966) and
Chung (1986), we know that nests are built on rocky cliff-
ledges and are constructed of dried sticks and grass. The
courtship behaviour is not known. Nests are described as
20-30 cm in diameter (Choi 1966).
The clutch size is reported as 4-6 eggs (Chung 1986). The
eggs are 'white and of prolonged ellipsoid shape with some
dotted markings' (H.G. Won 1966).
TAXONOMY
The Blackfaced Spoonbill has always been regarded as a
valid full species, because it is plainly distinct from the
sympatric Eurasian Spoonbill (Platalea leucorodid). However,
we believe that the question of the relatedness of this form
revolves not around the Eurasian Spoonbill but around the
Royal Spoonbill (Platalea regia) of Australia. It is possible
that the two forms are sufficiently closely related to be
considered races of one species (Kahl 1988a). If so, the
Blackfaced Spoonbill would be Platalea regia minor. The
possible affinities between Platalea minor and Platalea regia
were pointed out a hundred years ago by Seebohm
(1890:231).
It is commonly assumed that the Blackfaced Spoonbill,
sometimes called the Lesser Spoonbill, is a very small bird.
However, it is small only in relation to the Eurasian Spoon-
bill. There is in fact considerable overlap in size (see Appen-
dix) between Blackfaced and Royal Spoonbills.
As shown in the excellent colour-plate in Ogilvie-Grant
(1889:48, pl.l)t the soft-part coloration on the faces of the
Blackfaced and Royal Spoonbills is strikingly similar. It
appears that the Blackfaced Spoonbill usually has more
yellow under the eye, whereas the Royal Spoonbill usually
has more yellow over the eye. The Blackfaced Spoonbill also
appears to lack the red triangle at the top of the forehead.
The yellow eye-spots and red forehead triangles of living
I This plate is apparently not in all copies of The Ibis.
266 Blackfaced Spoonbill
spoonbills fade quickly after death, and both appear as dull
yellowish tan patches in preserved specimens.
It is sometimes claimed that the Blackfaced and Royal
Spoonbills can be separated by the extent and shape of the
bare black skin under the throat. Of course, this is not a
mark that can be used easily in the field. From our admit-
tedly limited experience with museum specimens, we feel
that this character is probably much too variable to be of
any use in separating the two populations (pers. obs.
M.P.K.). However, as with most aspects of the biology of
this spoonbill, further study on this point is desirable.
A bit of trivia regarding this species, to show the genetic
relatedness of spoonbills and ibises, is that a female Platalea
minor hybridized successfully with a male Threskiornis melano-
cephalus in the Berlin Zoo. The hybrid male offspring had the
head feathering of the mother spoonbill and the bill of the
father ibis. The hybrid later paired with a female Platalea
ajaja and produced a number of young (O. Heinroth 1905,
A.P. Gray 1958:5).
CONSERVATION
Considering the small total population, this bird must be
regarded as one of the more seriously endangered in the
world. According to H.G. Won (1966) the 'breeding
grounds on Ai island and other islands off the west coast of
North Korea have long been proclaimed [a] sea-bird's sanc-
tuary and guard boats patrol the territorial waters . . . Dur-
ing the breeding period fishing boats and other ships are not
allowed to approach the breeding islands. Moreover, land-
ing on the islands and collection of eggs there are strictly
prohibited.' According to Chung (1986) and Sonobe and
Izawa (1987:37) these areas are still protected by the North
Korean government. This is an extremely encouraging situ-
ation.
A sizeable portion of the population winters at the Mai Po
marshes, Hong Kong, where it currently receives strict
protection. The Mai Po area is well guarded because it is
not only a nature reserve but is also adjacent to the Hong
Kong/China international border. Steps should be taken to
assure the integrity of this area—and the ecological health of
the adjacent Deep Bay feeding areas—after 1997, when
Hong Kong merges with the People's Republic of China.
The main wintering grounds in Taiwan are said to be
under a threat of reclamation for industrial development or
an airport (Lucia Liu Severinghaus pers. comm. 1990). And
the Hong River estuary, Vietnam, has been greatly reduced
in the last few years to make shrimp-ponds and rice-
paddies. There remain only 6000 ha of mudflats, mangroves
and islets in the main delta area. Spoonbills are sometimes
taken for food in nets or snares in Vietnam. There are plans
to establish a nature reserve in the Hong River estuary
(D.A. Scott pers. comm. 1990), a conservation measure to
be applauded.
Any wetland that regularly holds more than 1 % of the
world population of any species of waterfowl is considered to
be of international importance under the Ramsar Conven-
tion. In the case of the Blackfaced Spoonbill, therefore, any
wetland holding 3 or more individuals is internationally
important (P. Kennerley pers. comm. 1990). At present,
however, not a single wetland regularly visited by Black-
faced Spoonbills has been proposed for inclusion under the
Ramsar Convention.
What losses may occur during migration across China,
and in other wintering areas along the South China Sea, is
unknown. However, direct persecution of migrating or win-
tering spoonbills is nearly certain in this densely populated
part of the world.
It is important that surveys be conducted to find addi-
tional nesting, migratory and wintering sites, especially in
China. The protection of all wintering and staging areas in
China, Korea, Taiwan and Vietnam would be highly desir-
able, as would the protection of additional breeding sites in
China, should they be found.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 325.
 African
Spoonbill
Platalea alba Scopoli

Platalea alba Scopoli, 1786, Deliciae Florae Fauna Insubricae, 2, p. 92; based on 'La Spatule blanche
de L'Isle de Luc.on' of Sonnerat, 1776, Voyage Nouvelle Guinee, p. 89, pi. 51: Luzon, Philippines
(error), Gape of Good Hope

Other names:
Red-legged Spoonbill (English); Spatule d'Afrique (French); Lepelaar (Afrikaans); Apamo (Luo);
Da-dosa (Mandingo), iNkenkane, isiXulamasele (Zulu); Sotrotsoina (Antakara, Sakalava/
Madagascar)

IDENTIFICATION
Standing 75-90 cm tall, the African Spoonbill is a medium-sized, rather stocky, bird with all-white
plumage, a grey spoon-shaped bill and red face.
In the adult the predominantly grey bill is edged on its upper surface with red; the lower surface is
grey and black, sometimes with yellow edges and spots. In addition to the face, the forehead, lores,
upper throat, legs and feet are red. The iris is pearly white to pale blue or pale grey.
When seen together, the sexes are distinguishable by their relative sizes. They look alike, but the male
is slightly larger and has a proportionately longer, heavier and slightly more decurved bill (Kahl 1983).
Changes at the start of breeding are subtle. The soft parts become a brighter red, the plumage a more
immaculate white. A fluffy and luxuriant crest develops, with loose plumes, rather than the lanceolate
ones of the Eurasian and Royal spoonbills. The crest becomes slightly tinged with pale cream colour at
the start of the breeding season, which may be from oil-gland secretions. The iris becomes a bright pale
blue, and the bill is paler bluish grey with brighter red edges (Kahl 1983).
At hatching the skin is pinkish and covered with white down; the bill is pink to orange and the legs
are pink. A description of nestlings at various stages of developement is given by Whitelaw (1968). The
spatulate tip on the bill begins to develop when the chick is about 11 days old. Immature birds are
generally similar to adults, but can be distinguished by the following differences: the feathers extend a
bit further forward on the face and forehead; the h^ad is streaked with blackish brown, and the crest is
small or non-existent; the tips of the primaries and some under wing-coverts are sooty black; the bill is
yellowish to horn-coloured; and the legs and feet are sooty grey to black.
The African Spoonbill is usually silent. Rather soft, nasal 'Kor, kor, kor' notes, having a crow- or
gull-like quality, are heard in flight, especially as a flock is taking off or preparing to land, and at
breeding colonies during displays (FJ. Jackson 1938:90). Also rather loud wing-noise occurs during
landing, take-off and in flight. It is especially noticeable around the breeding colony (Kahl 1983). Faint
rattling sounds, audible at close range during copulation, are produced by the male beating his bill back
and forth against the bill of the female; however, the soft, leathery bills of spoonbills are incapable of
producing the loud clattering sounds given by some species of storks. Aggressive interactions at the nest
elicit a hoarse 'Kwaark' (Whitelaw 1968). 'Yip-yip' calls are given during copulation.
The bird flies with its head and neck extended and legs trailing. When in a flock, spoonbills usually

fly in single file or diagonal line formation. Their flapping
rate averages 228/min (n = 25) (pers. obs. M.P.K.).
In the northern extreme of its range this species may,
occasionally, coexist with the rarer Eurasian Spoonbill.
However, the African Spoonbill is slightly larger, has a bare
red face, grey (rather than black) bill, and red (rather than
black) legs. Although this species can be confused with
other wading birds at a distance, its heavier stature and
outstretched neck in flight distinguish it from the white
herons. The straight, spoon-shaped bill separates the spoon-
bill from the Sacred Ibis.
DISTRIBUTION AND POPULATION
This is the characteristic spoonbill of Africa, occurring from
southern Mauritania, Senegal, central Sudan and Ethiopia
(from about 17°N) south to Cape Province, South Africa,
and in Madagascar.
Migratory patterns are poorly known. It seems likely that
they are mostly nomadic wanderings, in response to local
rainfall irregularities, rather than true seasonal migrations.
None the less, numbers fluctuate seasonally: African Spoon-
bills are more common in winter than summer in parts of
southern Africa (Brown et al. 1982:208). Birds nesting in
Sudan are known to migrate north to Khartoum, Rahad
and Kassala in the late dry season (G. Nikolaus in Luthin
1984a). One bird ringed in Transvaal, South Africa, was
later recovered several hundred kilometres north in Zambia;
other ringed birds have been recovered as far as 680 km
from their colony of origin (McLachlan and Liversidge
1970:57). A young spoonbill ringed at the Nata Delta, in
northern Botswana, was later recovered in the Orange Free
State, South Africa (Tree 1978).
No estimates of the African Spoonbill's population size
are available. It is widespread but with a patchy distri-
bution, and varies from uncommon to common (Brown et
African Spoonbill 269
al. 1982:208). The one known breeding colony in southern
Sudan had 50-100 pairs in the mid-1980s (G. Nikolaus in
Luthin 1984a), and the Garsen colony, in eastern Kenya,
had over 100 pairs in January 1983 (Coverdale et al. 1983).
This species has apparently increased and extended its
breeding range southwestward in Cape Province, South
Africa, since the 1950s (PJ. Wilson 1957, Blaker 1967,
Neame 1968b, Siegfried 1968), possibly because of the in-
crease in farm-ponds in the area.
ECOLOGY
The African Spoonbill occurs in many aquatic habitats,
including shallow lakes, rivers, marshes, artificial ponds,
and along the coast in estuaries and coastal lagoons.
It feeds by Walking Slowly through shallow water,
Sweeping the bill from side to side. Prey is captured in the
'spoon' of the bill and swallowed with a backward toss of the
head. Occasionally it dashes about frantically, in pursuit of
fish or other aquatic prey that has been visually detected.
In Zululand, Natal, South Africa, spoonbills have been
observed feeding behind a wading hippopotamus in shallow
water (Pooley 1967a). They also feed in association with
herons (J. Reynolds 1965, Connor 1979). In Zululand,
Pooley (1968) observed a spoonbill following a Great White
Egret (Egretta alba), 'with his bill almost between the heron's
legs, apparently feeding off small insects and crustaceans
disturbed by the heron as it stalked through the
water... the heron flew off several times, but each time the
spoonbill followed, until the heron eventually turned and
pecked the spoonbill to drive it away5.
The diet consists primarily of small fish, aquatic inverte-
brates and, occasionally, locusts (McLachlan and Liver-
sidge 1970:57). Three stomachs collected by Rand
(.1936:336) in Madagascar contained 'twenty small fish and
six small crayfish, one large water beetle and many small
crayfish, one large water beetle'.
The African Spoonbill is gregarious when feeding and is
usually found in small flocks. Usually, it is a rather shy bird,
but sometimes it becomes quite tame at nesting areas after
the eggs are laid. Often the spoonbill will rest on the shore of
feeding areas or in nearby trees, standing on one leg with its
head rotated to the rear and bill nestled in the feathers of the
upper back.
BREEDING
This species is gregarious in its nesting behaviour. It breeds
in colonies of up to 250 nests, often in association with, but
somewhat segregated from, other waterbirds, such as ibises,
herons, storks, cormorants and darters (Brown et al.
1982:208, Kahl 1983). African Spoonbills usually nest over
shallow water, in trees (e.g. Acacia spp.), bushes and
reedbeds (Phragmites australis). Whitelaw (1968) found them
nesting on an island of reeds in a sewage lagoon. They
sometimes nest on the ground on rocky islets (Bannerman
1930:125, pers. obs. J.A.H.) or on rocky ledges (Quickel-
berge 1972).
Breeding seasons are variable throughout the species'
270 African Spoonbill
extensive range in Africa. African Spoonbills nest mainly in
the dry season from West Africa to the Sudan, in the rainy
season in East and Central Africa, and mixed in both wet
and dry seasons in South Africa (Brown et al. 1982:209,
Maclean 1985:76). Nesting is also variable from year to
year, being suspended in a site when the rains do not occur
(FJ. Jackson 1938:90).
Courtship displays of the African Spoonbill have been
described by Kahl (1983). During pair formation, unmated
males establish themselves at potential nest-sites, and
unmated females approach them. Males initially attack and
drive away any other spoonbill that comes near. Through
persistent and repeated approaches, accompanied by Dis-
play Flights and Bowing displays, a female is eventually
accepted at the nest-site by a male. During Display Flights,
the woofing flight-noise is particularly loud and rasping.
A pair is established when mutual Greetings and other
pairing displays are performed, and copulations follow. The
Greeting consists of raising and opening the bill and fully
erecting the crest, while uttering a series of loud calls;
sometimes the wings are spread or flapped slowly. Soon
after pairing, birds engage in several displays that appear to
help cement the pair-bond. The series Display Shake/
Display Preen/Bill Popping occurs in a fixed sequence. In
the Head Quiver, the erect head feathers shake slightly.
Birds also stand side by side on the nest and engage in
mutual Head Shaking, in which the bill is pointed vertically
downward and the head is shaken in a 'no' motion. In
Display Sleeping, birds spend long periods with their bills
tucked under the feathers of their back, as if sleeping, but
keep their eyes open.
During the Copulation display, one bird (probably the
male) gives short 'Yip, yip, yip' calls. Copulations last about
10 s and may occur as often as every 5-10 min.
Pairs cooperate to defend the nest-site against intrusion
by other spoonbills with Sparring and, sometimes, with
overt attack. In the Sparring display, the bird erects its
crest, gapes its bill and makes biting movements at the
opponent. The opponent may respond with a similar action.
Allopreening by mated birds is frequent, most often per-
formed by the female. They commonly preen each other at
the same time, and most often direct their attentions to the
feathers of the head and neck.
Both sexes build the nest, the male flying off to collect
most of the material; addition of material to the nest con-
tinues throughout the breeding season. A flat platform is
constructed with small sticks, twigs, or reeds and is some-
times lined with dry grass. Five nests in South Africa aver-
aged 48 X 37 cm and were 6-11 cm deep (Whitelaw 1968).
The clutch size is 2-4 eggs. Mean clutches were 2.8 (47
clutches in East Africa; Brown et al. 1982:209), and 2.6 (303
clutches in South Africa; Maclean, 1985:76). Eggs are dull
white to pale buff, blotched with various amounts of reddish
brown or dark brown. Incubation, by both sexes, begins
after the second or third egg is laid. Females are reputed to
incubate more by day, males more by night. In exposed
sites, incubating birds sit facing into the wind. The incu-
bation period varies from 25 to 29 days; the average is
26 days (Brown et al. 1982:209).
Nestlings are constantly guarded by one or the other of
their parents until about 16 days old. Feeding, by both
parents, is by regurgitation, the nestling placing its head
well inside the throat of the adult. At a South African colony
of 12 nests, parental feedings were nearly twice as common
in the late afternoon (15.00-17.30 h) than they were in the
morning or at midday. By 21 days of age, young begin to
wander away from the nest on foot or in short flights, and by
about 35 days they are able to fly out of the colony; however,
they are still fed by their parents until at least 46 days of age
(Whitelaw 1968).
Breeding success is generally less than one young reared
per pair (Brown et al. 1982:209). In a South African colony,
59.5% of the eggs hatched, and 49.1% of those young
survived to fledging, giving an overall survival rate of 34%
and 0.82 young per nest (Whitelaw 1968). Nestlings are
sometimes taken by birds of prey: Brown (1970:73) stated 'I
have seen [an African Fish Eagle, Haliaeetus vocifer] visit a
Spoonbill colony repeatedly till it had practically wiped out
every brood.'
TAXONOMY
We agree with Snow (1978:42) that the African Spoonbill is
closely allied with Platalea leucorodia, Platalea regia and Plata-
lea minor, and these may be considered to form a 'species-
group'. However, Platalea alba is distinct enough from the
other three, not to be included with them in a defined
superspecies.
CONSERVATION
As is the case for many African species, this spoonbill faces
no apparent major problems. Although it may be affected
adversely in some areas by the drainage of wetlands, the
expansion of cultivation and ponds may be having positive
effects in other areas. Eggs or nestlings are sometimes taken
by humans for food. African Spoonbills have been bred in
captivity (Wylie 1982).
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 326.
 Yellowbilled
Spoonbill
Platalea flavipes Gould

Platalea flavipes Gould, 1837, Proc. Zool. Soc., pt.v, p. 106: New South Wales, Australia

Other names:
Yellowlegged Spoonbill (English)

IDENTIFICATION
The Yellowbilled Spoonbill, standing 76-92 cm tall, with a wing-span of about 140 cm, is a rather stout,
whitish waterbird with a long neck and legs, and a flattened yellow bill.
The adult's entire plumage is a very pale off-white. The bill and legs are dull straw-yellow to grey,
and the bare skin of the face is pale yellow. The iris is a pale greyish yellow.
The male has a noticeably larger body than the female, as well as a longer bill and legs. Of 5 adult
males and 9 adult females examined in museums, the average culmen length of females was 85.8% that
of males (18.64 vs. 21.72 cm) and the average tarsus length of females was 90.4% that of males (11.88
vs. 13.14cm) (Kahl 1988a).
At the approach of the breeding season, adults moult into a cleaner-looking, very pale yellowish white
plumage. A ruff of stiffened, spiky, pale straw-coloured feathers develops on the lower foreneck. More
than 100 feathers comprise a typical ruff and the longest ones are 6.5-7.5 cm in length. Several long,
filamentous black or dark grey inner-secondaries contrast with the white feathers of the lower back. Also
during breeding, the bare skin of the face becomes a pale cyanotic bluish yellow, with areas of pinkish
blue. A thin line, 2-4 mm thick, of black skin surrounds the bare face at the junction with the feathered
head and neck. The iris ranges from a pale greyish yellow to a pale blue, and the eye is surrounded by a
narrow ring of pink skin. A crescent-shaped spot of dark red skin, variable in shape but usually about
9 X 25 mm in size, appears on the base of the bill in front of each eye. The legs become a brighter
greyish green-yellow (for additional details and coloured photographs see Kahl 1988a).
Hatchlings have pinkish skin and are covered with a sparse white to light grey down; their thick,
blunt bill is pinkish yellow. Juveniles appear similar to non-breeding adults, except that they have sooty
tips on some primaries, shorter apricot-coloured bills, a darker brown iris, and duskier grey to olive-
grey legs. The dark tips to the primaries are sometimes retained by birds that otherwise appear to be
fully adult; these may be older subadults.
The Yellowbilled Spoonbill is usually silent, except at the nest where a variety of wheezing, coughing
and hissing vocalizations are given during displays (Kahl 1988a). The voice was also described as a
'feeble reedy grunt' (Pizzey 1980:64).
This species flies with alternate shallow flaps and glides, with the neck and legs extended. The
flapping rate averages 218/min (n = 23) (pers. obs. M.P.K.).
This species is easily distinguished from the white herons by its stockier build, spatulate bill, and
outstretched neck in flight. It often occurs in the same habitat with the slightly smaller Royal Spoonbill,
from which it is distinguished by the pale yellow, as opposed to black, bill, face, and legs.

DISTRIBUTION AND POPULATION
This species occurs primarily on the Australian mainland.
Its main breeding area is in southeastern Australia (Blakers
et al. 1984:67). It has occurred as a vagrant on Lord Howe
Island, King Island, Kangaroo Island, Tasmania (all Aus-
tralian territory), and New Zealand (Rangaunu Bay, Kai-
taia, on North Island; Billing 1977).
The Yellowbilled Spoonbill is a nomadic bird, wandering
according to rainfall and local conditions of flooding.
Little is known of specific population numbers. This
spoonbill may be considered common but seldom abundant.
However, the species seems to be holding its own in most
areas. A maximum of 250 birds was recorded at Lake
Cowal, NSW, at a time when the number of Royal Spoon-
bills was 100, and Strawnecked Ibises 20000 (Vestjens
1975b).
ECOLOGY
The Yellowbilled Spoonbill is found more or less wherever
there is water. It uses various types of shallow water, includ-
ing farm-ponds, borders of lakes, swamps and rivers. It is
seldom found in tidal areas but does occasionally frequent
saltpans and saline swamps. It is found on small, isolated
waters more often than is the Royal Spoonbill.
The species is most often seen feeding in loosely associ-
ated pairs, and sometimes in small groups. It may also feed
in mixed-species aggregations, with herons, ibises and other
spoonbills. It seems to be intimidated by the somewhat
more aggressive Royal Spoonbill. Foraging usually takes
place in water less than 40 cm deep. The feeding areas at
Lake Cowal were often within 300-500 m of the nesting site
(Vestjens 1975b, pers. obs. M.P.K.).
As is the case for the other spoonbills, foraging is mainly
tactile. The partly open bill is placed in shallow water and,
in Head Sweeping, the bill is moved slowly or rapidly from
side to side. When a food item is contacted by the 'spoon'
portion of the bill, it is grasped, the head raised and the item
swallowed with a quick backward toss of the head. Birds
also feed in a more directed way by Intensive Search, in
which the bill is moved rapidly from side to side in short
Yellowbilled Spoonbill 273
strokes (Vestjens 1975b). It also uses Probing to explore
under plants and Grabbing to pick up a visually located
prey item.
Their Feeding takes place both in the day and at night,
although we found foraging activity to be low during the
middle of very hot days (pers. obs. M.P.K.). Two Yellow-
billed Spoonbills, watched continuously for 8 h by Vestjens
(1975b), fed for about 7 h and waded 12 km while feeding.
The Yellowbilled Spoonbill has a longer bill, with a
narrower spoon than the Royal Spoonbill (Vestjens 1975b).
Such differences in bill-shape and slower feeding move-
ments result in different types of prey being taken by the two
spoonbills in the same habitat.
The diet is mainly aquatic insects (Notonectidae, Corixi-
dae), but also crustaceans, small fish (e.g. the introduced
Gambusia affinis) and freshwater snails. Vestjens (1975b)
examined a total of 28 Yellowbilled Spoonbill stomachs. He
found they ate four times as many crustaceans and insects as
fish, and in doing so consumed far fewer fish than did Royal
Spoonbills feeding in the same habitat.
At Lake Cowal, Vestjens (1975b) observed that Yellow-
billed Spoonbills captured about 1.3 items per minute of
foraging during the non-breeding season (April-August)
and about 7.1 items/min in the same habitat during the
breeding months (September-March). A total of 792 items
were found in one bird's stomach.
Often this spoonbill rests on the shore or in dead trees,
with the neck retracted or with the head rotated and the bill
buried in the feathers of the upper back. It has the repu-
tation of being shy, distrustful and difficult to approach.
Some adult birds at the nest allow a close approach,
whereas others are extremely wary.
BREEDING
Yellowbilled Spoonbills usually nest singly, although some-
times within 30-75 m of another pair. Colonial nesting
seems to be exceptional; however, AJ. North (1913-14:17)
reported 40-50 nests of this species, plus many cormorant
nests, in one large red gum tree. Other waterbirds, such as
Little Pied Cormorants (Phalacrocorax melanoleucos), often
build their nests near a spoonbill's.
The nest-platform is constructed of sturdy sticks, usually
over water, on horizontal limbs or in forks of trees or, less
commonly, in bushes or rushes. They usually nest higher
than the Royal Spoonbill. Of the 28 Yellowbilled Spoonbill
nests found during the 1984-85 breeding season at Lake
Cowal, 27 were in river red gum (Eucalyptus camaldulensis)
and 1 in a river cooba tree (Acacia stenophylla}\ all the trees
were standing in shallow water. These 28 nests varied from
2.5 to 10.0m (average = 5.34 m) above the surface of
the water (pers. obs. M.P.K.). Yellowbilled Spoonbills
occasionally take over old nests of other large birds, such as
the Wedgetailed Eagle (Aquila audax), for use as their own
(pers. obs. M.P.K.).
The pair-formation process is poorly known, because
birds often arrive at the nest site already paired. It is
possible that the pair-bond may last for more than a single
season (Kahl 1988a). Because pair-formation apparently
274 Yellowbilled Spoonbill
occurs elsewhere and nests are widely spaced, there is less
social interaction, and this species is less demonstrative at
the nest than other, more colonial, spoonbills.
During the Greeting, shown between mates at the nest,
the feathers of the head, neck, neck-ruff and scapulars are
strongly erected. In this display, each bird emits a series of
4—10 explosive coughing hisses (Kahl 1988a). Birds often
perform mutual Head Shaking and Stick Shaking displays
at the nest with their mates, especially early in the breeding
season.
As noted above, this species is generally rather shy
around the nest. When approached, the bird on the nest
adopts the Erect Posture and gives 'hoarse coughing hisses'
in a series, 0.3-1.0 s long and 1-15 s apart (Kahl 1988a).
The clutch-size is 2—4, perhaps rarely 5. The eggs are
symmetrical, elongate oval in shape and dull, matt white.
Incubation is by both parents. The exact incubation period
is unknown but probably lasts about 24—26 days.
Nestlings use the Begging display, with up and down
movements of the head and vocalizations, to solicit food
from their parents. In very small young, the call is a weak
chittering, high-pitched trill; as they grow older, the call
becomes deeper and lower-pitched, beginning to resemble
some calls given by adults (Kahl 1988a). Young are fed by
regurgitation by both parents. During the 1984-85 season at
Lake Cowal, a total of 23 nestling feedings averaged 5.4 s in
duration (range = 4.1-7.2s). A brood was usually fed be-
tween 11 and 14 times by a parent, and most of the feedings
took place within 10-30 min after the parent returned from
a foraging trip (pers. obs. M.P.K.).
Nestlings leave the nest at about 4 weeks and roam over
nearby bushes and trees; they are able to fly at about
7 weeks (Frith 1976:91). If forced to leave their nest-tree
before they are able to fly, nestlings are able to swim
strongly.
The birds appear to suffer predation at the nest site. On
23 October 1984 an Australian Raven (Corvus coronoides) was
seen taking a 1-week-old nestling from a Yellowbilled
Spoonbill nest at Lake Cowal (pers. obs. M.P.K.). And, in
the same area on 19 October 1984, a 2 m carpet python
(Morelia spilotes variegata) was observed ascending to a Yel-
lowbilled Spoonbill nest containing eggs. Vigilance and
aggression by the attending parents were apparently suc-
cessful in this case, for the snake was gone the next morning
and the adults continued with normal incubation (Kahl
1988a).
TAXONOMY
The Yellowbilled Spoonbill has sometimes been placed in a
monotypic genus, Platibis. However, we agree with Amadon
and Woolfenden (1952:6) that all the spoonbills are similar
enough to be contained within a single genus, Platalea.
Certain morphological features, such as the spiky neck-ruff,
show some affinities with the Roseate Spoonbill, Platalea
ajaja. And the blackish, filamentous inner secondary plumes
that develop during the breeding season are reminiscent of
the ibises of the genus Threskiornis.
CONSERVATION
This species seems to be doing well, and no specific conser-
vation problems are apparent. Some shooting and nest-
robbing, for zoo collections, occurs sporadically at places
such as Lake Cowal, NSW, but overall losses do not appear
to be serious.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 327.
 Roseate
Spoonbill
Platalea ajaja Linnaeus

Platalea ajaja Linnaeus, 1758, Syst. Nat., ed. 10, p. 140; based chiefly on 'Ajaia Brasiliensibus' of
Marcgrave, 1648, Hist. Rerum Nat. Brasiliae, p. 204: 'in America australi' = Rio Sao Francisco,
eastern Brazil, ex Marcgrave, designated by Berlepsch, 1908, Novit. Zool., 15, p. 301

Other names:
Pink Spoonbill, Rosy Spoonbill, American Spoonbill, Pink Curlew, Flame Bird, Pink, Banjo-bill,
Flamingo (English); Sevilla [Cuba], Chocolatera [Yucatan] Cuchareta, Cucharon [= large spoon],
Cucharera, Chuchara, Espatula rosada, Flamenco, Flamenco espatula, Garza paleta, Garza
colorada, Garza cuchara, Garza rosada, Ganso cucharon, Pato rosado, Penitente, Pico de cuchara
rosada, Planeta, Plateada, Sidra (Spanish); Spatule, Spatule rouge (French/Creole); Rode Lepelaar
(Dutch/Netherlands Antilles); Lepelbek (Dutch/Surinam); Colherado, Colhereiro (Portugese/
Brazil); Ayaya (Brazilian Indian)

IDENTIFICATION
The Roseate Spoonbill is a stunningly pink wading bird, with the distinctive spatulate bill characteristic
of spoonbills. It is a medium-sized, rather stocky, bird standing 75-80 cm tall, with a wing-span to 120-
133 cm and weighing about 1.2-1.75 kg. It is the only spoonbill with a brilliantly coloured plumage.
The adult's wings, abdomen and tail-coverts are bright pink to carmine. The neck, upper back and
upper breast are white. The tail is a surprising tawny-buff to orange. The naked head is greenish, whilst
the legs are magenta-ruby, blackish at the joints and toes.
The sexes are similar in plumage but dimorphic in size. The male is somewhat larger in body and
bill-length. These differences are apparent when a pair is together.
During the breeding season all the colours brighten. The lesser wing-coverts and upper and lower
tail-coverts become a brilliant carmine, whereas the wings and abdomen become a brighter pink. A
patch of stiff, recurved feathers is present on the upper breast and lower foreneck; these are coloured
magenta-ruby to carmine. A yellow wash develops on the side, adjacent to the bend of the wing.
The soft-part coloration also intensifies. The bill is greyish green to greyish tan with mottled areas of
green, black and yellow. The bare skin of the head assumes a pale green to golden-buff colour. A black
area of skin develops around the ear-openings, at the edge of the facial feathering, and across the back of
the neck (Palmer 1962:534). The black area on the head is individually variable in size and shape (A.
Sprunt IV pers. comm. 1990). The iris becomes cherry-red to scarlet.
During courtship, some individuals develop bright saffron-orange gular-sacs and orange rings around
their eyes. We have seen this in captive birds (from Surinam) at Vogelpark, Walsrode, Germany, as
they were courting and preparing to nest. This coloration was first reported by Audubon (1840) from
the southern USA and subsequently from South America by other first-hand observers (W.E.D. Scott
and R.B. Sharpe 1912, Hudson 1920:125, Wetmore 1926:66). There seems to be a dearth of first-hand
reports of this colour change in North American birds, and some accounts may have been secondarily
derived (Baird et al. 1884:103, Oberholser and Kincaid 1974:133). However, A. Sprunt IV (pers.

comm. 1990) states that it does occur in Florida birds, that
it is of short duration and indicates 'the highest of high'
colours. Additional information is desirable on the gular-sac
and colour changes of various populations.
At hatching, the nestling weighs about 50 g and is covered
with a sparse, short, white down. The skin, including the
bill and feet, is a rich salmon-pink (see photo in R.P. Allen
1962; Bent 1926:16, R.P. Allen 1942:119, D.H. White et al.
1982). The skin of a newly hatched spoonbill is not blackish
as stated in Palmer (1962:534). At about 2 days, the legs
and feet begin to darken, and by fledging they are a dark
grey-brown. By about 7 days, the downy coat becomes
longer, thicker and woollier. Except for the developing
flight-feathers, the body remains largely covered in down
until about 21 days (D.H. White et al. 1982).
The bill remains basically flesh-coloured until fledging.
As in all spoonbills, the bill is initially tubular. The spoon
on the end begins to develop at about 9 days. Near fledging
(39 days), it is only 67% the length of the adults (D.H.
White etal. 1982).
Large nestlings are faintly pink and have almost fully
feathered faces and heads (W.B. Robertson pers. comm.
1990). The plumage of juveniles is mostly white, with a
slight and variable suffusion of pink on the tail and under
the wings. The head and face remain more fully feathered
than in adults. The iris is dark brown to black. There are
dusky tips to the wing-coverts and primaries. The bill is a
pale dirty yellow to dull green, and the legs and feet are dark
Roseate Spoonbill 277
grey-brown. As birds mature, the amount of pink in the
plumage increases. And the head loses feathers, until the full
adult plumage is reached at about 3 years. [There is a fuller
discussion of plumaged succession in R.P. Allen (1942:119-
121,123); however, adults may lose the bright carmine in
their wings and tail during the non-breeding season, and
this would alter Allen's sequence.]
'When disturbed or alarmed, [it utters] a low 'Huh-huh-
huh-huh, repeated rapidly, without change in pitch or vol-
ume' (Palmer 1962:535). Like the American White Ibis, it
makes a soft grunting 'Uh-uh-uh' note while feeding conten-
tedly. Low clucking calls are heard during change-over at
the nest and in threat displays. In giving all these calls, the
head is raised and the bill opened. We have heard a 'gut-
tural croak' —more guttural and more voiced than in other
species of spoonbills —during courtship displays in captive
birds, which may be the calls that R.P. Allen (1942:86)
referred to as 'low, cackling and clucking sounds.' Soft bill-
rattling is sometimes heard during courtship activities and
displays.
Probably referring of a colony at feeding time, Chapman
(in Bent 1926:17) described the nestling's Begging call as 'a
chorus of tremulous, trilling whistles.' We have found the
whistling, cheeping calls of nestlings to be quite distinctive,
but reminiscent of ibises (pers. obs. J.A.K.).
The Roseate Spoonbill has a strong direct flight, often in
flocks that arrange themselves in clusters, diagonal lines or
wedge-shaped formations. It flies with its neck outstretched,
flaps alternating with short glides. Large, but still partly
dependent young, follow adults in flight while Begging for
food (R.P. Allen 1962). The flapping rate has, apparently,
not been recorded. A variation of the normal flight pattern
was described by Ogden (1976): 'birds made a short up-
glide, with their backs arched and wings stiff and angled
down, before resuming normal flight.'
Bright pink adults could be confused with flamingos and
the scarlet phase of the American White Ibis. The Carib-
bean Flamingo (Phoenicopterus ruber ruber) is taller and thin-
ner. The American White Ibis is smaller. The spoonbill is
easily distinguished from both by its characteristic bill
shape, but many reports of the flamingos outside their
normal range actually refer to Roseate Spoonbills. Pale
juveniles might be mistaken for an egret or the white phase
of the American White Ibis. Again, the bill shape is diagnos-
tic.
DISTRIBUTION AND POPULATION
The Roseate Spoonbill's overall breeding range can be de-
fined by the Neotropics. It nests in the southern USA
(mainly near the coast in Texas, southwestern Louisiana,
southern Florida), the Bahamas (Great Inagua), Cuba, Isle
of Pines, Hispaniola and adjacent islands, Mexico (outside
the central plateau region), Central America (although
breeding records are lacking for some countries), northern
and eastern Colombia, coastal and inland Venezuela and
the Guianas, southern Brazil (infrequently reported from
the main part of the Amazon basin), Ecuador, Peru (mostly
east of the Andes), eastern Bolivia, Chile (no recent nest-
278 Roseate Spoonbill
ing), Paraguay, Uruguay, Argentina (more common in the
north, south sparingly to Buenos Aires Province) (Bond
1961:41-12, Narosky 1978, Vigil 1973, Short 1975, Blake
1977:203, J. Steinbacher 1979, American Ornithologists'
Union 1983:58, R.B. Smith and D.R. Breininger 1988).
The species is a casual visitor or accidental over a much
greater range and well into the temperate areas of North
and South America, including various parts of the northern
and western USA, Bimini and elsewhere in the Bahamas,
Jamaica, Puerto Rico, the Lesser Antilles, Trinidad, Aruba,
Bonaire, coastal Ecuador, southern Patagonia and the Falk-
land Islands (Baird et al. 1884, Bent 1926:22-23, Bond
1961:41-42, Herklots 1961:41, Voous 1983:60, J. Stein-
bacher 1979, American Ornithologists' Union 1983:58, Raf-
faele 1983, Luthin 1984a).
The species undertakes dispersals and migrations in vari-
ous parts of its range. Fledglings as well as adults from
Florida disperse rapidly northward from their breeding sites
after leaving the nest, with juveniles going as far as 400 km
(Bent 1926:21, W.B. Robertson et al. 1983). Unmarked
juveniles have been found even further northward (e.g. to
Tennessee and South Carolina coast) (W.B. Robertson
pers. comm. 1990). Individuals from Texas disperse north-
ward as well (Oberholser and Kincaid 1974:132; Sutton
1967:45^46). In Brazil, they spread out widely after nesting
is over (Belton 1984).
There are not enough recoveries of marked spoonbills
beyond the first year to demonstrate conclusively the
seasonal patterns of movement. R.P. Allen (1942 and in
Palmer 1962:536) believed that migratory exchange took
place between Florida and Cuba, but his view was based on
very limited information. Birds move out of Louisiana and
much of Texas in winter, returning in February-April
(Oberholser and Kincaid 1974:132). However, spoonbills
also winter in both Texas and Florida. Although numbers
have increased recently in both Florida and Texas/
Louisiana, there seems to be little interchange between the
two populations (W.B. Robertson et al. 1983). Clearly,
more marking is necessary before the pattern of movement
is clear.
Spoonbills appear to disperse within South America as
well. Although present in Rio Grande do Sul, Brazil,
throughout the year, they are scarcer in winter, when they
are more restricted to the coast (Belton 1984).
The Roseate Spoonbill appears to be a common bird
throughout the core of its range in South America, being
particularly abundant in the wet savannas of Brazil, the
Chaco and Mato Grosso (R.P. Allen 1942:30).
This species has always been less abundant at the periph-
ery of its range, in the southern USA and northern Argen-
tina. The current population trend is an overall increase in
the USA. The trend was generally downward from 1850-
1920; thereafter, recovery was fairly rapid on the Texas
coast and slower in southern Florida. In the summer of
1941, R.P. Allen (1942:29-30) estimated that the total US
population was 5698 birds (513 in Florida, 150 in Louisiana,
5035 in Texas), 35% of which were breeding adults. By
1976, it was estimated that there were over 6400 nesting
birds in this area (2400 in Texas, 2600 in Louisiana, 1400 in
Florida) (Kushlan and White 1977, Blacklock et al. 1978,
Custer et al. 1980, Portnoy 1981).
Complete censuses have not been carried out elsewhere,
but some numbers are available. These include 3230 pairs
on the Gulf coast of Mexico in April-May 1971 (Sprunt and
Knoder 1980), 600 pairs on Isla Rajaros and Isla San Pablo
in Costa Rica, and 10 pairs on the Surinam coast (Luthin
1984a). Today few are seen in Belize, and the colony there is
apparently no longer extant (Luthin 1984a, D.S. Wood and
R.C. Leberman 1987). A 1964 nesting colony in Honduras,
near Cauquira, numbered c. 300 pairs (Monroe 1968:58).
The species is uncommon in Panama, although it was
reported as breeding there until 1930 (Ridgely 1976:50).
Nearly 1000 birds were counted in aerial surveys in Vene-
zuela (Luthin 1984a), but they were not breeding at the
time of the surveys.
This species is rare on Hispaniola (Wetmore and Swales
1931, de Dod 1978), but it is common in Cuba, nesting on
the Isle of Pines and along the mangrove coast of the main
island (Garrido and Garcia 1975:31). It apparently does not
nest on the other Antilles.
ECOLOGY
Roseate Spoonbills feed in various shallow, relatively open,
aquatic sites. They utilize heavily saline, salt, brackish and
freshwater areas. Over most of the range, these include
pools, lagoons, marshes, flooded pastures, stock-ponds and
rice-paddies. They also forage in tidal ponds, mangrove
pools and streams, mudflats, and estuaries. Feeding areas
may be as far as 20-35 km from the roosting or breeding site
(J.C. Lewis, 1983, pers. obs. J.A.K.). Although Roseate
Spoonbills are usually found in the lowlands, a wandering
bird was seen at about 4400 m in the Peruvian Andes (Stott
1959). Feeding sites are usually rather shallow, with maxi-
mum foraging depths in Florida Bay being about 20cm
(Powell 1987).
Although they will feed singly, Roseate Spoonbills are
very gregarious, foraging in small or large flocks with other
spoonbills, herons, ibises or storks. They often join feeding
flocks, and we have attracted them into a feeding site using
decoys (Kushlan 1977e). They appear to feed cooperatively
with other spoonbills, and other species sometimes use them
as 'beaters', gaining a foraging advantage by following them
around (M.L. Morrison et al. 1978, J.K. Russell 1978).
As with the other spoonbills, food detection and capture
are primarily by feeling rather than sight. They typically
feed by Head Sweeping, with the bill placed in the water
between vertical and nearly horizontally forward. Bob Allen
(1947:162) described it graphically: 'The broad, spatulate-
shaped bill is swept from side to side in wide semi-circles,
the mandibles slightly parted, the tips digging gingerly into
the surface film of the soft bottom, beneath waters that are
nearly always opaque. Delicate, sensitive to the small wrig-
glings and the darting, skittish movements offish or prawn a
quarter of an inch in length or of insects of even lesser
dimensions, this keen, responsive instrument must serve as
both eyes and hands.'
The entire head and neck, or just the bill, is scythed

laterally until a prey item is encountered. Then the bill is
clamped shut, raised and the food swallowed with a back-
ward jerk of the head. Spoonbills may manipulate large or
awkward prey by shaking, biting or beating them for many
minutes. They mostly wade in shallow water, Walking
Slowly, and moving along with the flock. However, we have
seen them Running after prey that they see near the surface.
When feeding on active prey, such as killifishes, birds 'will
sometimes run from side to side and flap their wings, leap-
ing ahead in order to intercept a school of minnows', but
when feeding on slower prey, such as prawns, the pace is
more leisurely (R.P. Allen 1942:93). They will Grab at prey
located visually, and they also use Probing and Intensive
Sweeping to obtain prey in localized areas.
We have seen birds feeding in water past their chests,
with the entire head and neck immersed (see also photo in
Heinrichs 1986), but more often they forage in shallower
water.
Feeding can take place at any time of the day or night. In
tidal areas, feeding activity peaks at low tide (R.P. Allen
1942:93, J.C. Lewis 1983:2). In many areas the birds feed
frequently, or even largely, at night (R.P. Allen 1947, Fried-
mann and Smith 1950, Powell 1987). Many years ago John
James Audubon (1840, and in Bent, 1926:19) stated that
spoonbills 'are as noctural as the night heron'. Although this
is a bit exaggerated, we have seen them feeding actively at
night many times, in Florida Bay on the falling tide. R.P.
Allen (1957:16-17) suggested that at night many of their
prey items are more active and easier to catch. However, in
Florida Bay, spoonbills fly out of their colonies and roosts in
the morning to feed in mangroves and freshwater marshes
during the day, retiring to the roost at night (pers. obs.
J.A.K., W.B. Robertson pers. comm. 1990, P. Bjork pers.
comm. 1990). Especially at the end of the nesting season, we
have seen juveniles feeding together away from the adults.
The diet consists almost entirely of small aquatic organ-
isms, such as small fish (e.g. Cyprinodon variegatus, Gambusia
qffinis, Fundulus sp.), crustaceans (shrimps, prawns, crayfish,
crabs), insects (mainly Coleoptera and Odonata), occasion-
ally molluscs (snails, including periwinkles, slugs) (Halli-
nan 1924, Bent 1926:19, Palmer 1962:540-541, Haversch-
midt 1968:30). Plant material may also be ingested; this
seems usually to be done accidentally, along with animal
prey, but it is sometimes taken intentionally (R.P. Russell
1982).
Three full stomachs obtained in Florida during
February-March, contained fish (81%), water beetles
(11%), shrimp (5%), plant material (3%), and traces of
gastropods and aquatic bugs. One of those stomachs con-
tained a total of 246 small fish, plus other miscellaneous
items (C. Cottam and P. Knappen 1939). R.P. Allen
(1942:102) gives figures for stomach contents (by volume)
as: fish 62%, Crustacea 20%, insects 14%, vegetation 3%,
molluscs a trace. At least for the US population, immature
sheepshead minnows (Cyprinodon variegatus) are 'probably
the most important single species in the diet of the Spoon-
bill' (R.P. Allen 1942:95). More recent analyses of nestlings'
stomach contents in Florida Bay show about 85% fish and
15% palaemonid-shrimp (R. Bjork pers. comm. 1990).
Roseate Spoonbill 279
Most prey eaten is small (usually <30 mm), but captives
were able to swallow pieces offish as large as one's fist (R.P.
Allen 1942:101).
There are occasional reports of predation on spoonbills,
mainly the young, by birds of prey such as Peregrine Fal-
cons (Falco peregrinus) (E.S. Clark 1981) and Bald Eagles
(Haliaeetus leucocephalus) (Robertson pers. comm. 1975). Rac-
coons (Procyon lotor) and Great-tailed Crackles (Quiscalus
mexicanus) often eat the eggs (R.P. Allen 1942:59-60). In
fact, we find it likely that the distribution of nesting sites in
Florida Bay reflect their inaccessibility to raccoons. We
have seen eagles and vultures taking eggs and young from
the nest, from which the parents were absent. Hurricanes
(W.B. Robertson and H.B. Muller 1961), droughts, and
winter storms cause some mortality as well. Man is the chief
enemy (see Conservation). Ectoparasitic infestations some-
times result in a high mortality of nestlings and desertion by
the parents (R.P. Allen 1942:88). In Texas, birds have been
observed leaving the water where they have been feeding to
defecate (McDaniel et al. 1963); this may help to reduce the
cycling of parasitic infections.
BREEDING
The Roseate Spoonbill usually nests on islands away from
ground predators, in colonies of up to 500 or more pairs.
(R.P. Allen 1942:89, W.B. Robertson et al. 1983). In Brazil,
it nests on Ilha do Governado, at the gates of the city of Rio
de Janeiro (Sick 1985). In most areas, such as the Venezue-
lan llanos, it typically nests in the company of various
herons, ibises, cormorants and storks. Spoonbills are some-
times dominated by herons at the nest-site (R.P. Allen
1942:89).
The nesting season varies geographically, as would be
expected for such a widespread species.
Depending on the region, nests are built in bushes, low
trees —such as mangroves, Rhizophora mangle, and Avicennia
germinans in Florida—and sometimes in taller trees (up to
20-25 m high). Spoonbills also nest in rushes or, rarely, on
the ground in secure sites (J.W. Harrison 1978). In Texas,
where larger trees are not available, spoonbills use low
vegetation on dredged-material islands (R.P. Allen 1942:50,
D.H. White et al. 1982, J.C. Lewis 1983). In general, the
preferred vegetation height for nesting is greater at main-
land than at island sites, possibly because of decreased wind
and increased need for protection from terrestrial predators
on the mainland (J.C. Lewis 1983). R.P. Allen (1942:50) felt
that a suitable feeding area near the nest is essential for just-
fledged young to use for several weeks before they join
adults at more distant feeding grounds.
The nest is often described as 'frail'. We have found,
however, that it is a rather sturdy platform of sticks and
twigs, often lined with finer twigs, grasses, leaves, bark or
moss. At one Texas colony nests were on average 55 cm
(range 43-71 cm) wide and 10-13 cm deep (D.H. White et
al. 1982). R.P. Allen (1942:84) found nests in Texas 'more
substantial in their construction' than nests in Florida.
Birds arrive at breeding areas in flocks, presumably
280 Roseate Spoonbill
unpaired. Bob Allen (1942:73-74) described 'up flights' and
'sky gazing' as typical behaviours seen in the pre-pairing
stage. During 'up flights', flocks suddenly take wing en masse,
circle around, and then land again. In 'sky gazing' a group
of standing adult spoonbills react to another flying over by
extending their necks to full length, pointing their bills
skyward and appearing to gaze upward for a few seconds.
These two behaviour patterns seem not to have been
reported in other species of spoonbills. However, the 'up
flights' might possibly represent a group version of the
Display Flight, described for lone African, Royal and
Yellowbilled Spoonbills (Kahl 1983, 1988a).
Pairing birds also engage in the Display Preen. This
display of the Roseate Spoonbill, as witnessed in courting
captive birds at Vogelpark, Walsrode, Germany, was quite
different from the homologous displays of the other spoon-
bills so far studied (details in Kahl 1983, 1988a). In the
Roseate, the head is shaken in a very loose-jointed way, with
the tip of the bill describing a 'figure-eight'. Then there is a
quick, loose-jointed preen of the upper wing-coverts, the
primaries or the neck-ruff, which is erected. During court-
ship, some soft bill-rattling is heard, especially during the
Display Preen. Potential mates bite repeatedly at each
other's bills when excited. Displaying and courting birds
often fly and land with the tongue-bone extended, causing
the throat-skin (or gular-sac) to be conspicuously exposed
(pers.obs. M.P.K.).
Copulations, once begun, are frequent. In one pair the
first successful copulation was observed 14 days after pair-
formation (R.P. Allen 1942:89). In one period of 2 h 40 min,
a pair copulated three times and the male brought 11 twigs
to the nest (Palmer 1962:538). An apparently successful
copulation lasted 20-25 s (R.P. Allen 1942:84).
Other breeding displays described by R.P. Allen (1942)
and Palmer (1962:538) appear to be generally similar to
those observed in other species of spoonbills (Kahl 1983,
1988a). However, further detailed studies are needed to
fully describe the display repertoire of this species.
Both sexes cooperate to build the nest, with the male
collecting most of the material and the female doing most of
the building. They may use the nests of earlier-nesting ibises
(Weller 1967). There is apparently an increase in nest-
building activity just before the eggs hatch (R.P. Allen
1942:88).
Eggs vary from ovate to elliptic-ovate or even elongate-
ovate (Bent 1926:16). They are white, or nearly white, with
no gloss, and with spots and small blotches of various
shades of brown.
The clutch size is 1-5, usually 2 or 3. R.P. Allen (1942:34)
gives the average clutch size as 2.7. In a Texas colony, 87
nests had an average of 2.64 eggs per clutch on 27 May,
when egg-laying was complete (Palmer 1962:538). In 154
marked nests over a 3 year period in Texas, clutch size
varied significantly from year to year but averaged 3.0
(D.W. White et al. 1982). One female laid her first egg
6 days after the first observed copulation (R.P. Allen
1942:89). Eggs are usually laid every other day.
Incubation, by both sexes, begins the day after the first
egg is laid and lasts from 22 to 24 days, probably most often
22 days (R.P. Allen 1942:88, Palmer 1962:540, D.H. White
etal. 1982).
R.P. Allen (1942:34) stated that breeding adults are quite
nervous and 'will desert a nest that is disturbed'. Care must
be taken, as with other spoonbill species, in approaching the
nests for observation or photography, especially before eggs
have been laid. However, comparisons between intensively
studied and relatively undisturbed colonies (D.H. White et
al. 1982) show that careful observations can be done with-
out causing significant mortality.
Both parents feed their young by regurgitation. Young
put their bills in the side of the adult's bill and down into the
throat to obtain the meal. Parental feedings last about 10s
(F.M. Chapman 1914, and in Bent 1926:17). Young leave
the nest at about 4—5 weeks (R. Bjork pers. comm. 1990).
They remain nearby and are fed by parents until at least 8
weeks.
Nesting success varies widely, depending on environmen-
tal conditions. In some years success is good, whereas in
others most nests may be lost. R.P. Allen (1942:34) reports
that
In some seasons a colony appears to hatch and rear all
eggs and young; in other seasons I have recorded nest
mortality as high as 98.5%. The average nest mortality
for Florida Bay for the five seasons from 1936-37 to
1940-41, inclusive, is estimated at about 35%. In
Texas the average nest mortality over a six-year period
has been about 22% . . . the average number of young
produced annually per pair is about 1.8 in Florida Bay;
2.1 in Texas.
At a recent Texas colony, D.H. White et al. (1982) found
that among 154 nests in 3 seasons, 73% of the eggs laid were
hatched and 50% of the eggs laid were fledged, giving an
average production of 1.5 young per nest.
Roseate Spoonbills probably breed, at the earliest, in
their third year (R.P. Allen 1942:33). Although Allen stated
that 'only adults in full breeding plumage pair', Dunstan
(1976) observed successful breeding by adult spoonbills that
lacked the full breeding plumage. Pairing is probably for a
single breeding season only (R.P. Allen 1942:34).
TAXONOMY
The Roseate Spoonbill has sometimes been placed in a
monotypic genus, Ajaia, because of its distinctive plumage
and its simplified trachea (R.P. Allen 1942:126, Vestjens
1975a). However, we agree with Mayr and Short (1970) and
J. Steinbacher (1979) that it should be included within the
genus Platalea. Certain morphological features, such as the
spiky neck-ruff and lack of nuchal crest, show an affinity
with the Yellowbilled Spoonbill (Amadon and Woolfenden
1952). It may be more closely related to this species than to
spoonbills of the Eurasian/Royal/Blackfaced group.
CONSERVATION
The large and widespread populations of Roseate Spoon-
bills throughout tropical America have apparently

remained more or less stable. Based on its history, this
species appears to be rather resilient in recovering and
maintaining its population levels. It also may be more
resistant to the effects of pesticides than some other species
of waterbirds (Oberholser and Kincaid 1974:133, D.H.
White etal. 1982).
In North America and Cuba, however, the species was at
one point brought to the verge of extinction (Sprunt 1939,
R.P. Allen 1942, T. Barbour 1943:19, Oberholser and Kin-
caid 1974:133). At the low point, between 1890 and 1919,
there may have been only 20-25 breeding pairs left in the
USA. 'If a reservoir of supply had not existed in countries to
the south, it would probably have become extinct in the
United States before 1900' (R.P. Allen 1942:14).
The drastic and widespread decline of the US population
came during the plume-hunting period in the late 19th
century. Spoonbills were occasionally killed so that their
wings could be made into fans. However, because the
feather colour fades rapidly, the Roseate Spoonbill was not,
itself, hunted on a large scale for the millinery trade. Their
decline was probably brought about by the wanton disturb-
ance of breeding areas, also containing egrets that were
hunted on a large scale for their plumes. According to R.P.
Allen (1942:3) 'the shooting of egrets in colonies that these
birds shared with the "Pink Curlews" would have had an
effect on both species, even though no Spoonbills were
actually killed. Death of the egrets would, in my opinion,
coincide with desertion of the area by the Spoonbills.'
During the current century, populations in the southern
USA have shown a dramatic recovery, first on the Texas/
Louisiana coast and later, and to a lesser extent, in southern
Florida. Without a doubt much of that recovery was due to
increasingly stringent wildlife legislation and vigilant pro-
tection of breeding areas in sanctuaries, primarily those
administered by the National Audubon Society.
In Texas and Louisiana, recovery occurred between 1920
and the 1970s. The Roseate Spoonbill began to reappear as
a breeder on the Texas coast in the 1920s, and by 1976, it
was estimated that there were over 2400 nesting birds in
Texas and 2600 in southwestern Louisiana (Blacklock et al.
1978, Portnoy 1981).
Recovery in Florida came more slowly, possibly because
Florida Bay, the spoonbill's main nesting ground there, was
still too remote for adequate law enforcement. Spoonbills
were still being exploited, mainly for food, through the
1930s (Powell et al. 1989). Partial recovery did eventually
occur in Florida, and Robert Porter Allen (1962) could
Roseate Spoonbill 281
write: 'From a single tiny colony of 15 pairs in 1939, the
Florida nesting population increased to a peak of eight
colonies and more than 400 breeding birds in 1955-56'.
From 1955 to 1978 the nesting population in Florida Bay
roughly doubled every 10years (Powell et al. 1989). By
1978-79 there were approximately 1400 breeding pairs in
southern Florida (W.B. Robertson et al. 1983). They are
now breeding, in small numbers, as far north as Tampa Bay
(west coast), where they had not nested since 1912 (Dun-
stan 1976), and on Merritt Island (east coast) (W.B.
Robertson pers. comm. 1990). They also are expanding
their range from west to east in Louisiana (Portnoy 1981).
Surveys during 1984-87 suggested that the Florida Bay
breeding population dropped back from the 1978-79 high of
1254 pairs to approximately 450-590 pairs (Powell et al.
1989). At the same time, populations of other fish-eating
birds using Florida Bay have also suffered reproduction and
population decreases. These include the Osprey (Pandion
haliaetus} (Kushlan and Bass 1983), the Brown Pelican (Pele-
canus occidentalis] (Kushlan and Frohring 1985), and the
Great Blue Heron (Ardea herodias] (Powell 1983). These
simultaneous declines are probably due to a common cause,
the reduction of food availability (Kushlan and Frohring
1985).
The source of these problems may lie in disturbance to
the ecology of south Florida. Reduced bird populations stem
from decreased production of young in years past. This may
be due to inadequate food resources in Florida Bay, which
in turn may be caused by man-made changes in the fresh-
water run-off from the Everglades. Additionally, commer-
cial and residential development in the Florida Keys has
claimed wetlands that were once used by foraging spoonbills
(Powell etal. 1989).
The very high numbers (1254 pairs) in Florida Bay dur-
ing 1978-79 may just be an anomaly, reflecting the
unusually high recruitment of new birds from a very suc-
cessful season 3 years before; except for that one year, the
Florida Bay population has been at 550-750 pairs for the
past 15-20 years (A. Sprunt IV, pers. comm. 1990). Accord-
ing to W.B. Robertson (pers. comm. 1990), there has been a
considerably better nesting success in Florida Bay during
the past 2-3 years.
Note: Body measurements, egg measurements and nesting
season data for this species can be found in the Appendix on
page 328.
This page intentionally left blank
Appendix
 American Wood Stork
(Mycteria americana)

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

Male [3] 464-475 227-240 196-205 186-204 1664-1790 (1) Extremes 35 62.5-72.3 43.8-48.8 (1)
Male, average [3] 470 233 200 195 1727 — (1) Average 35 67.7 46.7 — (1)
Male [6] 453-492 215-235 194-213 155-166 — — (2) Extremes 40 60.8-73.0 34.3-54.9 — (2)
Male, average [6] 476 227 203 158 — (2) Average 40 67.9 46.0 — (2)
— _
Male [2] 455-479 221-244 — (5) Average 20 68.1 46.5 — (3)
— —
Male, average [2] 467 233 — — — — (5) Average — — — 83 (4)
Male [4] — — — — 3.05-3.30 (6) Extremes 64 62.0-73.0 43.0-51.0 — (5)
—
Male, average [4] — — — — 3.18 (6) Average 64 68.0 46.0 — (5)
— _
Male [2] — 205-230 197-205 — — (10) Calculated 64 — — 80 (5)
Male, average [9] — — — — 2.70 (7) Average — 67.8 46.0 — (6)
—
Male [?] 452-484 215-239 197-214 155-160 — _ (13)
Male, average [?] 468 227 208 158 _ (13) References: (1) Pers. obs. M.P.K. (2) Bent 1926:60. (3) Palmer 1962:513.
—
Male [1] — — — — 3.09 (1) (4) O. Heinroth 1922. (5) Schonwetter 1967. (6) Oberholser and Kincaid
—
Male [1] 225 — — 2.50 (10) 1974:125.
— —
Male [1] 493 235 208 155 — (12)
—
Fern. [6] 435-455 190-205 175-192 143-150 — (2)
—
Fern., average [6] 446 196 183 148 — (2)
—
Fern. [?] 422-464 192-208 183-202 132-155 —
— (13) EGG-LAYING
Fern., average [?] 443 202 193 146 —
— (13)
Fern. 460 205 — — — (5)
[1]
—
Fem. [1] — — — — — 2.05 (7) Place Months
Fem. [1] — — — — — 2.78 (8)
Fem. — — 2.40 (9) S Florida (formerly) Nov-Dec (a few Jan+)
[1] — — —
Fem. [1] — 205 185 — — 2.15 (10) S Florida (currently) Jan-Mar (a few Nov-Dec)
Fem. — 200 180 — — (10) Central/N Florida Feb-May
[1]
—
Fem. — — — 2.80 (11) Georgia, South Carolina Apr-Jun
[1] — —
Both [?] 443-488 — 175-213 — — — (3) SE Texas May-Jul
Both, average [?] — — — — 1650 — (3) Mexico (Tabasco, Campeche) Jan-Mar
_
Both [?] — — — 1550-1650 (4) Honduras Jan-Feb
—
Both, average [?] — — — — 1650 — (13) Costa Rica Dec-Jan
Venezuela (Llanos) Sep-Dec
Longevity in wild = 11 years, 8 months (pers. obs. M.P.K.; from USFWS Banding Office). Surinam (coastal) Sep-Oct
Longevity in captivity = 30-1- years (M. Davis pers. comm., 16 July 59). Brazil (Matto Grosso) Jul-Sep
Body temperature = 40.65°C (n = 66) (Kahl 1963b). Argentina (E central) Oct-Nov
Breathing rate = 11-12/min (Kahl 1963b).
References: (1) Pers. obs. M.P.K. (2) Blake 1977:187. (3) Eaton 1910:244. (4) Forbush and May 1955:41. (5)
Palmer 1962:510. (6) Hartman 1955. (7) Hartman 1961:41. (8) Haverschmidt 1968:26. (9) O. Heinroth 1922.
(10) B.T. Thomas 1985:923. (11) Tashian 1953. (12) Sharpe 1898:322. (13) Oberholser and Kincaid 1974:124-
125.

284
 Milky Stork
(Mycteria cinered)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Fern. [1] 435 201 200 145 (1)

Fern. [1] 450 195 190 153 — — (5)
Both [?] 4.4.4 494
194-238 188-225 156 — — (3)
Both, average [?] 470 220 188 160 — — (1)
p 438 203 188 170 — — (2)
IU
p 495 270 223 165 — — (4)
[1]
p 500 275 225 165 — — (5)
[1]

References: (1) Kuroda 1936:539. (2) Pers. obs. M.P.K. (Malaysian Game Dept./25 Jan 68). (3) H.C.
Robinson and F.N. Chasen 1936:206. (4) Sharpe 1898:326. (5) Delacour and Jabouille 1931:93.

EGG M E A S U R E M E N T S

Length Width Weight

No.. (mm) (mm) (g) Reference

Extremes 44 59.0-74.5 43.0-48.0 (1)

Average 44 66.3 45.9 — (1)
Calculated 44 — — 77 (2)
Extremes 24 62.0-68.2 43.3-47.9 — (3)
Average 24 64.8 45.4 — (3)
Extremes 3 62.7-64.1 43.9-45.7 — (3)
Average 3 63.3 45.1 — (3)
Extremes 3 66.0-70.0 45.0-46.0 — (4)
Average 3 68.3 45.7 — (4)

References: (1) Hoogerwerf 1949b:19. (2) Schonwetter 1967. (3) Helle-

brekers and Hoogerwerf 1967:13. (4) Pers. obs. M.P.K. (Mus. Zool., Bogor/8
Mar 68).

EGG-LAYING

Place Months

WJava Mar—Aug
SE Sumatra Jun-Aug
W Malaysia Aug

285
 Yellowbilled Stork
(Mycteria ibis)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Fern. [1] 425 200 200 175 (10)

Fern. [1] — — — — — 1.19 (11)
Fern. [1] — — — — — 2.33 (11)
Both [?] 455-513 205-242 197-229 168-183 1500-1650 — (1)
Both [?] 450-515 205-260 190-230 160-190 — — (2)
Both [?] 450-475 208-229 200-221 175 — — (3)
Both [?] 455-513 205-242 197-229 168-183 — — (4)
Both [?] 450-500 205-230 190-230 160-180 1500 — (5)
Both [?] 490-526 — — — — — (6)
Both [?] — — — — 1500-1650 — (7)
Both [?] 455-517 203-243 195-229 168-190 1500-1650 — (8)
Both [?] 455-517 203-243 195-218 170-190 — — (9)
? [1] 500 245 225 183 — — (12)

Longevity in captivity = 19-1- years (K. Brouwer et al., pers. comm., 1990).
References: (1) Brown et al. 1982:173. (2) Schouteden 1954. (3) Heuglin 1869-73:1130. (4) Bannerman
1930:108. (5) Reichenow 1900-01:334. (6) Clancey 1964a:47. (7) Cramp 1977:322. (8) Maclean 1985:70. (9)
McLachlan and Liversidge 1970:50. (10) Andersson 1872:296. (11) Britton 1970b. (12) Sharpe 1898:328.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
West Africa: Gambia to N Nigeria Nov-Feb
Extremes 17 59.0-72.0 42.0-46.0 (1) Gambia, Senegal, Guinea-Bissau Nov-Feb
Average 17 66.0 44.0 — (1) Ghana Nov-Jan
Calculated 17 — — 71 (1) N, central Nigeria Aug-Jan
Extremes — 59.0-66.0 40.0-45.0 — (2) Chad Jun-Sep
Average — 61.3 42.0 — (2) S Sudan Oct-Nov
Extremes 15 62.1-71.1 42.3-47.6 — (3) S Somalia Oct
Average 15 66.7 45.3 — (3) W Kenya, Uganda Mar-Jun
Extremes 3 65.0-68.0 44.0-46.0 — (4) NW Kenya [L. Turkana] Aug-Sep
Average 3 66.0 45.0 77 (4) E Kenya Jan, May-Aug
Extremes 36 59.0-72.3 42.0-46.1 — (5) NW Tanzania Jan-Jul
Average 36 66.0 44.0 — (5) S Zaire Jul-Aug
Extremes 51 59.0-72.3 42.0-47.6 — (6) Zambia Feb-Sep
Average 51 66.2 44.4 — (6) Malawi Jul-Sep
Single — — — 62 (7) NE Botswana Mar, Aug-Sep
NE South Africa (Natal) May-Oct
References: (1) Schonwetter 1967. (2) Reichenow 1900-01:334. (3) W. Fraser
1971. (4) Mundy and Cook 1974. (5) Brown et al. 1982:175. (6) Maclean
1985:71. (7) Heinroth 1922.

286
 Painted Stork
(Mycteria leucocephald)

BODY MEASUREMENTS EGG MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

Male [1] 500 203 156 (7) Extremes 60 65.0-80.0 43.2-51.6 (i)
Male [1] 513 263 — — — — (9) Average 60 70.0 48.0 — (i)
Fern. [1] 488 275 240 150 — — (9) Calculated
Extremes
60 —
66.0-75.0
— 89 (i)
Fern. [1] 525 225 194 163 — — (7) 60 41.5-48.0 66-85 (2)
Fern. [1] 488 — 230 155 — — (7) Average 60 69.6 43.9 74 (2)
Fern. [1] 505 215 210 — — — (10) Extremes 27 64.5-73.8 43.8-49.5 — (3)
Both [?] 490-510 252-278 240-250 150-172 — — (2) Average 27 69.3 47.0 — (3)
Both [2] - — — — - 2.02-3.5 (3) Extremes 50 65.3-80.2 43.2-51.6 — (4)
Both [?] — — - 2.0-3.0 (4) Average 50 65.9 45.0 — (4)
—
Both [?] — — — 1500-1600 - (8) Extremes 3 67.6-68.2 47.3-48.5 — (8)
Both [?] 490-500 215-278 210-250 150-172 — — (10) Average 3 67.9 47.7 — (8)
Both [3] 450-510 — — — — — (10) Extremes 2 — 82-98 (5)
—
Both [6] 490-523 230-256 205-242 — — — (5) Average 50 69.5 49.0 — (6)
Both, average [6] 505 246 225 — — — (5) Average — 71.0 48.0 — (7)
Both [10] - — — — - 3.03-3.37 (6)
Both, average [10] - — — — - 3.18 (6) References: (1) Schonwetter, 1967. (2) Desai etal. 1977. (3) Hume and Gates
Both, average [?] 495 279 243 152 — — (1) 1890:223. (4) E.C.S. Baker 1935:451. (5) O. Heinroth 1922. (6) S. AH and S.D.
Ripley 1968:94. (7) Henry 1971:385. (8) Pers.obs.M.P.K. (N India/lOSep67).
Longevity in captivity = 28+ years (K. Brouwer et al. pers. comm. 1990).
References: (1) T.H. Shaw 1936:120. (2) E.G.S. Baker 1929:332. (3) S. AH and S.D. Ripley 1968:95. (4) O.
Heinroth 1922. (5) Abdulali 1968. (6) Desai et al. 1974. (7) Legge 1880:1100. (8) Tom Roberts pers. comm. (9)
Sharpe 1898: 325. (10) Delacour and Jabouille 1931:92.

EGG-LAYING

Place Months

Pakistan Jun-Nov
N India Jul-Oct
S India Nov-Mar
Sri Lanka Mar-Apr
Burma Aug-Nov
Thailand (Ban Yang, formerly) Jul
Kampuchea (Grand Lac) Jan-Feb

287
 Asian Openbill Stork
(Anastomus oscitans)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 388 153 158 160 (4)

Fern. [1] 400 160 150 155 — — (4)
Fern. [1] 385 — — — — — (5)
Fern. [1] 370 158 134 — 1190 — (6)
Both [?] 400-425 158-163 150-158 150-188 1475-1488 — (1)
Both [?] 392-408 153-162 140-150 183-217 — — (2)
Both, average [14] - 160 — — — — (3)

Longevity in captivity = 7+ years (K. Brouwer et al. pers. comm. 1990).

References: (1) Legge 1880:1103. (2) E.C.S. Baker 1929:333. (3) Kahl 1971c:27. (4) Sharpe, 1898:307. (5)
Delacour and Jabouille, 1931:94. (6) Pers. obs. M.P.K. (northern India/13 Sep 67).

EGG MEASUREMENTS

Length Width Weight

No. (mm) (mm) (g) Reference

Extremes 80 48.3-64.0 36.4-43.4 (1)

Average 80 57.8 41.1 — (1)
Average 40 55.0 37.3 — (2)
Extremes 100 48.3-63.1 36.4-45.2 — (3)
Average 100 57.9 41.2 — (3)
Extremes 57 50.0-63.0 37.0-45.5 — (2)
Average 57 56.0 40.0 — (2)
Extremes 30 51.0-62.0 36.4-44.2 — (4)
Average 30 58.0 41.5 — (4)
Calculated 30 — — 55 (4)
Extremes 2 55.8-59.7 40.0-41.2 — (6)
Single — 59.0 41.0 — (5)

References: (1) E.C.S. Baker 1929:334. (2) Hume and Gates 1890:225.
(3) E.C.S. Baker 1935:454. (4) Schonwetter 1967. (5) Herbert 1926. (6) Pers.
obs. M.P.K. (N India/10 Sep 67).

EGG-LAYING

Place Months

N India, Burma Jun-Sep

S India Nov—Mar
Sri Lanka Dec-Apr
Thailand Nov-Feb

288
 African Openbill Stork
(Anastomus lamelligerus]

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

A. I. lamelligerus Extremes 39 51.0-61.4 36.7-43.2 35-50 (i)

Male, average [16]
Male
—
432
172
194
—
160
—
—
—
—
—
—
(6) Average
Extremes
39
14
55.4
51.0-59.5
40.0
39.5-43.2
—
—
(i)
[1] (11) (2)
Male [1] — — — — — 1.25 (7) Average 14 54.7 40.5 c. 50 (2)
Fern. [1] — — — — — 1.0 (7) Extremes 13 53.5-55.5 36.7-41.5 — (3)
Fern. [1] — — — — — 1.14 (8) [sic]
Fem. [1] — — — _ — 1.26 (12) Average 13 55.7 39.9 — (3)
_
Fern. [1] — — — — 1.17 (12) Extremes 5 53.4-59.8 38.2-40.3 — (4)
Both [?] 400-420 160-180 140-160 180-210 1400 — (1) Average 5 56.2 39.3 — (4)
Both [?] 384-436 — — — — — (2) Extremes 12 52.0-60.5 36.5-40.5 — (5)
Both [?] 369-400 154-171 119-146 185-196 — — (3) Average 12 56.4 39.0 — (5)
Both [30] 370-433 153-194 133-165 165-206 1400 — W Extremes — 55.3-61.4 40.8-41.3 — (6)
Both [?] 400-440 — — — — — (5) Extremes 4 56.0-58.0 40.0-42.0 — (7)
Both [?] 390-435 — — — — — (10) Average 15 53.3 39.9 — (8)
? [1] 393 175 160 175 — — (9) Extremes 10 51.0-59.5 39.5-43.2 — (9)
Average 10 54.4 40.9 — (9)
A. I. madagascariensis Calculated 10 — — 50 (9)
Male [1] 400 153 133 — — — (11)
Fem. [1] 360 140 130 — — — (11) References: (1) Maclean 1985:68. (2) Brown etal. 1982:177. (3) Anthony and
Sherry 1980. (4) Benson and Pitman 1958. (5) Braine 1974. (6) Bouet 1955a.
References: (1) Reichenow 1900-01:336. (2) Macworth-Praed and Grant 1973:67. (3) Heuglin 1869-73: 1120. (7) F.J.Jackson 1938:75. (8) Heiden 1974. (9) Schonwetter 1967.
(4) Bannerman 1930:102. (5) FJ. Jackson 1938:73. (6) Kahl 1971c. (7) Brown et al. 1982:176. (8) Maclean
1985:68. (9) Sharp 1898: 309. (10) Clancey 1964a:45. (11) Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (12)
FitzPatrick Inst., Univ. of Cape Town (unpubl. data).

EGG-LAYING

Place Months

Chad Sep
Ethiopia (L. Tana) Aug-Sep
Kenya, Uganda Recorded all months
Tanzania Jan-Apr/Jul-Aug
Zambia Recorded all months
Zimbabwe Sep-Jun
NE Botswana Jan-Feb
N Namibia Jan-Mar
N South Africa Feb-Mar
Madagascar Sep-Dec

289
 Black Stork
( Ciconia nigra)

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Length Width Weight

Wing Culmen Tarsus Tail Wing-Span Weight No. (mm) (mm) (g) Reference
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference
Extremes 11 61.2-67.1 46.1-49.0 (1)
Male [1] 3.2 (15) Average 11 64.0 47.0 — (1)
Male [1] 592 188 213 260 - — (16) Extremes 84 60.3-74.3 45.2-51.7 — (2)
Male [1] 560 205 195 250 - — (17) Average 84 65.4 48.8 — (2)
Male [1] 538 193 205 200 - — (21) Extremes 47 60.0-74.0 45.0-56.0 — (3)
Fern. [1] 525 175 190 213 - — (21) Average 47 69.0 48.7 c. 86 (3)
Fern. [1] 559 185 205 237 - — (16) Extremes 21 63.0-74.0 45.0-56.0 — (4)
Both [?] 520-580 — — — — — (1) Average 21 69.4 48.8 — (4)
Both [?] 520-600 — — 230-250 1900 3.0 (2) Extremes 6 64.5-69.5 45.0-48.3 — (5)
Both [?] 520-550 170-195 190-205 250 - — (3) Average 6 66.7 47.3 — (5)
Both [9] 520-600 160-190 180-200 190-240 - — (4) Extremes 3 70.0-74.0 48.5-50.0 — (6)
Both [?] 530-570 180-195 185-195 230-250 - 3.0 (5) Average 3 71.3 49.2 — (6)
Both [?] 516-580 — — — — — (6) Extremes 3 68.5-69.5 48.5-49.0 — (7)
Both [?] 540-615 — — — — — (7) Average 3 68.8 48.8 — (7)
Both [?] 525-590 170-195 185-200 — — — (8) Extremes 3 68.0-73.0 47.8-48.0 — (8)
Both [?] 530-570 180-195 185-195 230-250 - — (9) Average 3 70.0 47.9 — (8)
Both [7] 520-575 175-194 182-207 220-245 — (10) Average — — — 83 (9)
Both [4] 540-585 193-198 198-204 245-264 - — (10) Average 50 65.5 49.0 — (10)
Both [?] 520-600 160-190 180-(280?) 190-240 1440-1550 c. 3.0 (11) Extremes — 60.0-74.3 44.0-54.7 86 (11)
Both [9] 520-600 160-190 180-200 — — 3.0 (12) Extremes 20 62.8-70.8 47.3-51.8 — (12)
Both [?] — — — - 1440-1550 — (13) Average 20 67.0 49.6 — (12)
Both [8] 520-580 150-190 175-200 — — — (14) Extremes 80 (60.0)-72.0 44.0-53.0 — (13)
Both, average [8] 539 — — — — c. 3.0 (14) Average 80 65.4 48.8 — (13)
Both [?] 520-600 160-195 180-200 190-250 3.0 (19) Calculated 80 — — 86 (13)
? [1] 550 185 — - 1550 3.0 (18)
p [1] — — — — — 2.59 (20) References: (1) Dementiev and Gladkov 1951:442. (2) Hartert 1912-21:1216.
p [1] — — — — — 2.25 (20) (3) Maclean 1985:66. (4) McLachlan and Liversidge 1970:53. (5) Priest 1933.
(6) P.R. Barnes 1950. (7) Ellenberger 1951. (8) Howells 1968. (9) O. Heinroth
Longevity in wild = 18 years (Bauer and Glutz von Blotzheim 1966:424). 1922. (10) Etchecopar and Hue 1978:73. (11) Schroder and Burmeister
Longevity in captivity = 30 years (Dresser 1871-81:316). 1974:31. (12) Adam Mrugasiewicz pers. comm. (6 April 1969). (13) Schon-
Longevity in captivity = 314- years (K. Brouwer et al. pers. comm. 1990). wetter 1967.
References: (1) Clancey 1964a:43. (2) Sokolowski 1968:146. (3) Reichenow 1900-01:347. (4) McLachlan and
Liversidge 1970:52. (5) O. Heinroth and M. Heinroth 1924-31:139. (6) Mackworth-Praed and Grant 1973:64.
(7) Meinertzhagen 1954:390. (8) Kozlova and Tugarinov 1947:263. (9) Hartert 1912-21:1216. (10) Bannerman EGG-LAYING
1930:97. (11) Brown et al. 1982:178. (12) G. Niethammer 1938:299. (13) Cramp 1977:323. (14) Dementiev and
Gladkov 1951:445. (15) S. Ali and S.D. Ripley 1968:104. (16) LaTouche 1931-34:440. (17) Taczanowski 1891-
93:975. (18) Paz 1987:33. (19) Schroder and Burmeister 1974:5-6. (20) FitzPatrick Inst., Univ. of Cape Town Place Months
(unpubl. data). (21) Sharpe, 1898:305.
Europe Apr-Jun
USSR, Mongolia, NE China, SE Korea Apr-May
Zambia May-Jun
Malawi Jul
Zimbabwe Jan-Sep (mainly May/Jun)
Mozambique Sep-Oct
Botswana Jun
N South Africa, Lesotho May-Sep (mainly Jun/Jul)
S South Africa (Cape Prov.) Aug-Sep

290
 Abdim's Stork
(Ciconia abdimii)

BODY M E A S U R E M E N T S EGG MEASUREMENTS

Length Width Weight

Wing Culmen Tarsus Tail Wing-Span Weight No. (mm) (mm) (g) Reference
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference
Extremes 41 56.2-64.1 40.1-45.8 (1)
Male [1] 438 110 120 180 (5) Average 41 59.6 43.3 — (1)
Fern. [1] 435 105 118 175 — — (5) Extremes 23 55.0-63.5 39.7-47.1 — (2)
Fern. [1] — — — — 1.08 (4) Average 23 58.3 42.5 — (2)
—
Both [22] 400-475 103-127 117-136 167-205 — (1) Calculated 23 — 58 (2)
— —
Both [?] 413-425 108-113 115-119 175 — — (2) Extremes 4 55.0-61.0 40.0-47.0 — (3)
Both [?] 430-460 100-120 110-130 170-200 — — (3) Extremes 37 54.6-64.8 40.8-45.3 — (4)
p [1] — — — — 0.98 (4) Average 37 60.4 43.1 — (4)
p —
[1] — — — — — 0.94 (4) Extremes 11 56.2-62.0 40.5-44.0 — (11)
9
[1] — — — — — 0.93 (4) Average 11 58.3 42.0 — (11)
Extremes — 51.0-56.0 39.1-41.1 — (5)
Longevity in captivity = 16 years 5+ months (Flower 1925:1399). Average — 52.1 (37.5?) — (5)
Longevity in captivity = 2 1 + years (K. Brouwer et al. pers. comm. 1990). Extremes — 50.0-58.0 40.0-42.0 — (6)
References: (1) Bannerman 1930:100. (2) Heuglin 1869-73:1105. (3) Reichenow 1900-01:344. (4) FitzPatrick Average — 59.6 43.3 — (7)
Inst. Univ. of Cape Town (unpubl. data). (5) Sharpe 1898:293. Singles 1 55.5 41.0 — (8)
Singles 1 58.0 41.5 — (8)
Singles 1 60.8 40.0 — (9)
Singles 1 60.3 39.6 — (9)
Singles 1 — — 60 (10)

References: (1) Serle 1943 in Brown et al. 1982:181. (2) Schonwetter 1967.
(3) Reichenow 1900-01:344. (4) Kahl 1971d. (5) Heuglin 1869-73:1107. (6)
Bouet 1955a. (7) Bannerman 1953:173. (8) Bannerman 1930:101. (9) Bigalke
1948. (10) Salvan 1967. (11) Duignan et al. 1988 (K. Brouwer pers. comm.
1990).

EGG-LAYING

Place Months

W Africa to Ethiopia, Somalia, N Kenya Feb-Jul (mainly Apr-May)

Equatorial Kenya and Uganda Jan-May
E Eritrea (winter rainfall area) Jan-Mar
SW Arabia, Yemen Mar-May

291
 Woollynecked Stork
(Ciconia episcopus}

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

C. e. episcopus C. e. microscelis
Male M 471-507 149-185 161-178 175-205 - - (1) Male [1] 500 163 175 180 - — (24)
Male, average W 488 162 170 192 - - (1) Fem. [1] - _ _ _ _ 1.59 (19)
Male [7] 471-523 — — _ _ _ (2) Fem. [1] 463 143 155 180 - — (24)
Male, average [7] 493 — — _ _ _ (2) Fem. [1] ~ _ _ _ _ 1.85 (23)
Male [8] — 143-165 — _ _ _ (3) Fem. [1] ~ _ _ _ _ 1.59 (23)
Male [7] 446-500 137-157 140-163 _ _ _ (21) Both [?] 454-491 _ _ _ _ _ — (14)
Male, average [7] 469 148 157 _ _ _ (21) Both [?] 413-438 117-131 129-144 175-179 - — (15)
Male [8] 440-500 155-171 — _ _ __ (25) Both [16] 440-485 135-168 140-171 180-214 - — (16)
_ _ _
Male, average [8] 478 163 (25) Both [?] 450-490 _ _ _ _ — (17)
— _ _ __
Male [1] 515 — — (9) Both [?] 440-470 130-140 145-150 170-180 - — (18)
Male [1] 519 — — — — — (9) p [1] 440 140 152 177 - — (20)
~\/f
Male i — — 1.93 /(10)
1 r\\
Male [1] — — — 2.5 (10) Longevity in captivity = 30+ years (K. Brouwer et al. pers. comm. 1990).
Male [1] 465 146 — 182 (13) References: (1) Riley 1925:29. (2) Abdulali 1968. (3) Medway and Wells 1976:94. (4) Meyer and Wiglesworth
Male [1] 464 137 — 193 - - (13) 1898:807. (5) Legge 1880:1119. (6) E.C.S. Baker 1929:324. (7) Hackisuka 1931-32:348. (8) Kuroda 1936:540.
Male [1] 497 145 160 169 - - (22) (9) S. Ali and S.D. Ripley 1968:99. (10) J.H. Riley 1938:36. (11) duPont 1971 :23. (12) Rabor 1977 :26. (13)
Male [1] 525 — — _ _ _ (22) Rensch 1931. (14) Mackworth-Praed and Grant 1970:53. (15) Heuglin 1869-73:1109. (16) Bannerman1930:98.
Fern. [3] 460-480 146-147 153-166 193-195 - - (1) (17) Clancey 1964a:45. (18) Reichenow 1900-01:348. (19) Britton 1970a. (20) McLachlan and Liversidge
Fern., average [3] 472 147 158 194 - - (1) 1970:51. (21) Pers. obs. M.P.K. (Zool. Mus., Bogor, Indonesia/8 Mar 68). (22) Delacour and Jabouille 1931:
Fern. [4] 450-516 — — — — — (2) 85-86. (23) FitzPatrick Inst., Univ. of Cape Town (unpubl. data). (24) Sharpe 1898:296. (25) Pers. obs.M.P.K.
Fern., average W 485 — — _ _ _ (2) (British Mus./28 Aug 70).
Fern. [2] — 148-152 — _ _ _ (3)
Fern. [5] 442-462 140-157 147-160 — — — (21)
Fern., average [5] 450 145 154 _ _ _ (21)
Fem. [10] 435-480 140-171 — _ _ __ (25)
Fern., average [10] 458 150 — —
_ —
_ —
_ (25)
Fem. [1] 490 — — (22)
Both [9] 440-490 125-169 — _ _ __ (25)
Both, average [9] 472 153 — — — — (25)
Both [5] 475-490 137-157 — _ _ _ (4)
_ _ _
Both average [5] 485 146 — (4)
Both [?] 450-515 — 150-175 175-205 1750-1850 2.05-2.79 (5)
Both [?] 444-497 145-168 152-180 199-225 - - (6)
Both [?] 470-508 141-145 157-179 183 - - (7)
Both [?] — 165-176 152-158 158-166 - - (8)
Both P] 444-525 145-168 152-190 169-190 - - (22)
? [1] 470 [131?] 148 193 - - (11)
P 450 153 155 173 - - (12)
[1]

292
Woollynecked Stork continued

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
C. e. episcopus
Extremes 3 58.8-62.3 45.0-47.5 (1) N India May-Sep
Average 3 60.8 46.2 — (1) S India Nov-Mar
Extremes 100 57.0-72.3 43.6-49.3 — (2) Sri Lanka Jan-Apr
Average 100 62.9 47.4 — (2) Burma Feb-Mar/May-Jul/Sep
Extremes 50 57.5-66.5 43.8-48.0 — (3) Malaysia (Tanjong Badak) Mar-Apr
Average 50 62.5 45.8 — (3) Philippines Jan
Extremes — 60.0-61.0 41.0-42.0 — W Indonesia (Flores) Feb-Mar/Jun/Aug/Nov
Extremes — 59.0-64.0 47.0-48.0 — (5)
Extremes 3 59.3-61.2 44.7-45.4 70-70 (6) C. e. microscelis
Average 3 60.4 45.0 70 (6) Liberia Sep-Oct
Extremes 3 60.0-61.3 41.2-42.3 — (7) Sierra Leone Jan-Feb
Average 3 60.7 41.9 — (7) N Nigeria Jan
Extremes 7 59.3-67.0 43.0-45.4 53-70 (9) Chad Jul-Sep
Average 7 62.0 44.4 59 (9) Sudan (Upper Nile) Get
Singles 1 67.0 43.0 55 (8) Ethiopia J«l
Singles 1 65.7 43.2 55 (8) Uganda Nov-Feb
Extremes 4 60.0-67.0 41.2-43.2 — (10) Zambia Aug-Sep
Average 4 63.5 42.4 — (10) Zimbabwe Aug-Nov
Calculated — — — 59 (10) N South Africa (Zululand, Natal) Sep-Nov
Extremes 120 57.0-72.0 43.6-49.3 — (11)
Average 120 63.0 47.3 — (11)
Calculated 120 — — 77 (11)

References: (1) Inglis 1898. (2) E.C.S. Baker 1935:443. (3) Hume and Gates
1890:269. (4) Bannerman 1953:172. (5) Bouet 1955a. (6) Anthony 1978b. (7)
Pitman 1931a. (8) Anthony 1977. (9) Maclean 1985:67. (10) Brown et al.
1982:182. (11) Schonwetter 1967.

293
 Storm's Stork
(Ciconia stormi)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 405 163 163 155 (1)

Male [1] 398 — — 136
_ -
_ -
_ (2)
Male [1] — 137 136 (3)
Fern. [1] 375 143 138 133 - (1)
Fern. [1] 390 — 138 143 - - (1)
Fern. [1] 404 — — 143 (2)
Fern. [1] 420 147 150 180 - - W
Both [5] 380-470 120-135 — _ _ _ (6)
Both, average [5] 402 131 — _ _ _ (6)
p [1] — 168 — _ _ _ (4)
P _ _ _
[1] 395 — 150 (5)
? [1] 395 — 150 — — — (5)

References: (1) Ogilvie-Grant 1903. (2) Mayr 1938. (3) Pers. obs. M.P.K. (Zool. Mus., Bogor, Indonesia/8
Mar 68). (4) Medway and Wells 1976:95. (5) Medway and Wells 1964:156. (6) Pers.obs. M.P.K. (British Mus./
28Aug70).

EGG M E A S U R E M E N T S

Length Width Weight

No. (mm) (mm) (g) Reference

Single 60.2 41.9 (1)

Reference: (1) F. Danielsen et al. pers. comm. 30 Apr 1990.

EGG-LAYING

Place Months

S Thailand Sep
SE Sumatra Apr-May

294
 Maguari Stork
(Ciconia maguari)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [8] 225-255 265-275

—
__
_
_
_
_
—
(i)
Male [6] 531-614 203-245 (2)
Male, average [6] 562 231 266 200 - — (2)
Male, average [9] — 241 — _ _ 4.2 (6)
Male [1] — 235 260 — — 4.0 (1)
Male [1] — 230 260 — — 4.2 (1)
Male [1] — 235 260 — — 4.25 (1)
Male [1] — 230 270 — — 4.35 (1)
Fern. [6] — 205-227 — — — — (1)
Fern. [5] 535-566 205-229 — — — — (2)
Fern., average [5] 553 219 — — — — (2)
Fern., agerage [5] — 213 — — — 3.8 (6)
Fern. [1] — 200 240 — — 3.4 (1)
Fern. [1] — 205 235 — — 3.5 (1)
6oth [2] 535-603 207-227 — 196-208 - — (3)
Both, average [2] 569 217 — 202 - — (3)
? [1] 590 230 280 250 - — (4)
P [1] 540 — — — — — (5)
Longevity in captivity = 20+ years (K. Brouwer et al. pers. comm. 1990).
References: (1) B.T. Thomas 1985:923. (2) Blake 1977:188. (3) Goodalletal. 1951:101. (4) Zotta and Fonseca
1937. (5) Chubb in Haverschmidt 1968:27. (6) C.E. King 1988:162.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
Venezuela (Llanos) May-Nov (mainly Jul-Sep)
Extremes 54 69.1-85.0 46.0-57.0 100-140 (i) E Brazil (Isla Mexiana) Aug-Sep
Average
Extremes
54
18
75.2
70.0-84.0
52.6
46.0-57.0
116
—
(i) NE Argentina (Corrientes Prov.)
E-central Argentina (Buenos Aires Prov.)
Jun-Aug
Jul-Oct
(2)
Average 18 75.1 53.4 — (2)
Calculated 18 — — 118 (2)
Single 1 77.4 56.2 — (3)
Single 1 76.3 56.2 — (3)
Single 1 75.6 51.9 — (4)
Single 1 74.0 52.0 — (5)
Single 1 70.4 50.0 — (6)
Single 1 71.5 51.5 — (6)
References: (1) B.T. Thomas 1984. (2) Schonwetter 1967. (3) Kahl 1971e. (4)
Gibson 1880. (5) Gibson 1919. (6) Haedo-Rossi 1969.

295
 White Stork
(Ciconia ciconid)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference
&
C. c. ciconia ~^ C. c. asiatica
Male [9] 530-630 150-190 195-240 215-240 — — (1) Male [14] 580-630 184-235 - (4)
Male [?] 600-620 180-200 — 240-260 — — (2) Male, average [14] 610 - — - - - (4)
Male [41] — — — — — 2.9-4.4 (3) Male [18] 581-615 188-223 213-247 - - - (22)
Male [9] 540-610 169-206 — — — — (4) Male, average [18] 596 204 234 - - - (22)
Male, average [9] 580 — — — — — (4) Male [1] 653 220 - - - - (17)
Male [18] 556-598 157-198 191-230 221-268 — — (22) Fern. [9] 548-596 178-196 211-234 - - - (22)
Male, average [18] 576 179 214 247 — — (22) Fern., average [9] 577 187 220 - - - (22)
Male, average [41] — — — — — 3.571 (3) Both [?] 590-615 195-215 - - - - (15)
Male, average [12] — — — — — 3.97 (5) Both [?] 600-620 198-222 - - - - (15)
Male [1] — — — — — 4.3 (14) Both [ 9 ] 550-640 (16)
Fern. [?] 530-590 140-170 — — — — (1) Both [14] 580-630 184-235 - - - - (13)
Fern. [?] 560-580 150-175 — — — — (2) Both [16] - 188-210 - - - - (18)
Fern.
1 [27] — — — — — 2.7-3.9 (3) Both, average [9] 589 - - - - - (16)
trem., average ro7i 0 OO^
o.azO /Q\
U'J W)

Fern [15] 543-582 155-180 184-211 218-256 — — (22) a

Longevity, oldest ringed bird = 29 years 3 months (Rydzewski 1973).
b
Fern., average [15] 558 164 197 237 — — (22) Longevity, free-flying semi-captive = 33 years 8 months (Bloesch 1986).
c
Fern., average [12] — — — — — 3.52 (5) Longevity in captivity = 33 years 3 months (Flower 1938).
d
Fern. [1] — — — — — 3.5 (11) Longevity in captivity = 35-1- years (K. Brouwer et al. pers. comm. 1990).
Both [?] — — — — 1550-1650 — (6) References: (1) Witherby et al. 1939:115. (2) Hartert 1912-21:1214. (3) Bauer and Glutz von Blotzheim
Both [6] 585-605 — — — — 3.5-4 (7) 1966:390. (4) Vaurie 1965:84. (5) Hornberger 1967:7. (6) Cramp 1977:328. (7) Dementiev and Gladkov
Both [6] 542-580 158-191 213-225 218-251 — — (8) 1951:451. (8) Cramp 1977:335. (9) Reichenow 1900-01:346. (10) O. Heinroth and M. Heinroth 1924-31:133.
Both [?] 550-600 175-200 190-220 210-240 — — (9) (11) O. Heinroth 1922:185-186. (12) Mell 1951:24. (13) J. Niethammer 1967. (14) Milstein 1966 .(15) Hartert
Both [9] 540-610 169-206 — — — — (13) 1912-21:1215. (16) Dementiev and Gladkov 1951:453. (17) Mendelssohn 1975a. (18) G. Niethammer
Both [?] — — — — up to 2000 — (19) 1972b:208. (19) Schierer 1972. (20) Grzimeketal. 1968:208. (21) Creutz 1988:8. (22) S.W7. Winter pers. comm.
Both [?] 530-630 — — — up to 2200 2.3-4.4 (20) 1990.
Both [?] — — — — — 2.6-4.5 (21)
Both, average [12] 565 176 220 227 — — (8)
Both, average [?] — — — — — 3.5-4 (10)
Both, average [?] — — — 'almost — (12)
— 2.5m'

296
White Stork continued

EGG MEASUREMENTS EGG-LAYING

Length Width Weight

No. (mm) (mm) (g) Reference Place Months

Extremes 150 65.0-81.5 46.5-56.0 (1) Spain, Portugal Feb-May

Average 150 73.0 51.8 — (1) N Europe Apr-Jun
Calculated — — — 110 (1) W and central Asia Apr-Jun
Extremes 103 65.6-81.5 46.5-55.7 — (2) NW Africa Feb-Apr
Average 103 73.2 51.8 — (2) S South Africa Sep-Nov
Extremes 27 68.6-77.6 49.0-55.9 95.5-129.3 (3)
Average 27 77.3 52.5 111.5 (3)
Extremes 20 65.6-81.5 46.0-55.7 — (3)
Average 20 70.7 50.7 — (3)
Extremes 80 61.6-78.6 47.0-54.5 — (3)
Average 80 71.3 51.5 — (3)
Extremes 16 71.0-78.2 47.9-57.0 — W
Average 16 73.8 53.8 — (4)
Extremes 120 65.5-81.5 46.5-55.7 — (5)
Average 120 73.2 [58.8?] — (5)
Extremes 140 64.0-79.0 48.0-53.2 — (6)
Extremes 13 — — 95.5-129.3 (7)
Average — — — 120 (8)
Extremes 37 66.6-88.6 48.5-57.1 — (10)
Average 37 73.2 52.3 — (10)
Single 1 — — 118 (9)

References: (1) Schonwetter 1967. (2) Witherby et al. 1939:114.

(3) Bauer and Glutz von Blotzheim 1966:403. (4) Dementiev and Gladkov
1951:449. (5) E.C.S. Baker 1929:322. (6) Haverschmidt 1949:40. (7) G.
Niethammer 1938:297. (8) O. Heinroth and M. Heinroth 1924-31:133. (9)
Mell, 1951:25. (10) Adam Mrugasiewicz pers. comm. 6 Apr 69.

297
 Oriental White Stork
(Ciconia boy eland)

BODY M E A S U R E M E N T S EGG MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex M (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference
Male, average [5] 266 301 (1) Extremes 10 68-77 53-58 (1)
Male [11] 616-670 235-279 256-300 245-307 — — (2) Average 10 75.0 54.5 __ (1)
Male, average [11] 654 257 277 271 _ — (2) Calculated 10 — — 124 (1)
Male [8] 580-678 223-276 280-315 250-285 - 4.5-5.5 (3) Extremes 24 72-80.3 51.5-58.3 — (2)
Male, average [8] 645 243 296 264 5.22 (3) Average 24 76.7 56.3 — (2)
Male [14] 592-680 207-282 250-310 — _ — (4) Extremes 4 76-79 58-60 (3)
Male, average [14] 640 256 — — — —
282 (4) Average 4 77.5 59 147.5 (3)
Male [2] — — — - 4.13-5.5 (4) Extremes 4 74-79 56-58 — (4)
—
Male, average [2] — — — — 4.82 (4) Average 4 77.3 56.9 — (4)
Male [2] 650-690 — 290-305 238-260 _ — (6) Extremes 4 78.1-81 57-62.3 — (5)
_ _
Male, average ?
670 260 298 249 (6) Average 4 79.6 58.4 — (5)
Male, average [9] — —
— _ 5.1
_ (8) Extremes 10 72.7-80.2 52.1-55.6 — (8)
—
Male, average [31] — 255 285 — (11) Average 10 78.9 54.6 — (8)
Male [16] — 226-276 — — — — (12) Extremes — 77-80 54-56 — (6)
Male, average [16] — 248 —
— — — (12) Average — 75.2 58.1 — (7)
Male [1] 665 — 265 245 — — (9) Average — 77.0 58.0 — (9)
Male [?]
Fern., average
[1] 590
—
227
239
241
271
223 —
_
—
_
(10)
[7] — (1) References: (1) Schonwetter 1967. (2) Winter 1982:78. (3) Fei et al. 1991.
Fern. [7] 598-655 222-288 241-266 216-284 — — (2) (4) Leonovich and Nikolyevsky 1976. (5) Pankin and Neufeldt 1976. (6) Austin
Fern., average [7] 615 243 253 242 _ _ (2) and Kuroda 1953:333. (7) E.C.S. Baker 1929:323. (8) S.V. Winter pers. comm.
Fern. [7] 540-658 210-245 245-278 220-265 - 3.7-5.0 (3) 1990. (9) Taczanowski 1891-93:974.
Fern., average [7] 615 224 265 252 - 4.35 (3)
Fern. [20] 576-635 203-248 215-282 — — — (4)
Fern., average [20] 606 228 255 — — — (4)
Fern. [4] — — - 2.89-4.5 (4)
— — EGG-LAYING
Fern., average [4] — —
— — 3.75 (4)
Fern. [3] 615-705 231-288 245-265 230-256 — — (6)
Fern., average [3] 653 255 252 241 _
— _
— (6) Months
Place
Fern., average [26] — 232 254 — (H)
Fern., average [8] — — — — - 4.1 (H) Mar
Korea (formerly)
Fern. [15] — 210-245 — — — — (12) Japan (formerly) Mar-May
Fern., average [15] — 224 — — — — (12) Apr-May
SE USSR/NE China
Both [?] 620-670 195-222 —
—
— — (5)
Both [8] 620-670 200-260 — — — — (7)

Longevity in captivity = 31 years (Schtiz 1965b).

Longevity in captivity = 48+ years (C.E. King and K. Brouwer et al. pers. comm. 1990).
Body temperature of large nestlings; 6 readings = 40.5-41.0°C (Fei et al. 1991).
References: (1) K. Archibald and B. Schmitt 1991. (2) S.W. Winter pers. comm. 1990. (3) K. Murata, unpubl.
data (via K. Archibald pers. comm.). (4) K. Archibald pers. comm. 1990. (5) S. Ali and S.D. Ripley 1968:102.
(6) Taczanowski 1891-93:974. (7) Hartert 1912-21:1215. (8) C.E. King pers. comm. 1990. (9) LaTouche 1931-
34:438. (10) T.H. Shaw 1936:121. (11) C.E. King 1988:162. (12) Murata et al. 1988.

298
 Blacknecked Stork
(Ephippiorhynchus asiaticus]

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 563 300 300 213 (3)

Fern. [1] 538 288 275 213 — — (3)
Both [4] 488-625 288-300 295-338 235-250 — — (1)
Both, average [4] 566 297 313 242 2125 — (1)
Both r?i 565-645 298-324 300-333 257-281 — _ (2)
p [i] 573 320 323 200 — — (4)
? [i] — — — — c. 2000 — (5)

Longevity in captivity = 34+ years (K. Brouwer et al. pers. comm. 1990).
References: (1) Legge 1880:1116. (2) E.C.S. Baker 1929:326. (3) Sharpe 1898:312. (4) Rand and Gilliard
1967:48. (5) Pizzey 1980:62.

EGG MEASUREMENTS

Length Width Weight

No. (mm) (mm) (g) Reference

Extremes 50 67.0-76.0 51.0-57.0 (i)

Average 50 72.0 53.7 — (i)
Calculated
Extremes
50
11
—
71.0-76.0
—
52.0-55.0
115 (i)
— (2)
Average 11 74.5 53.8 — (2)
Calculated 11 — — 120 (2)
Extremes 45 66.3-78.3 49.5-57.5 — (3)
Average 45 72.8 53.0 — (3)
Extremes 30 67.9-74.9 51.0-55.2 — (4)
Average 30 69.5 53.2 — (4)
Extremes p 70.3-71.9 52.4-54.0 — (5)
References: (1) Schonwetter 1967 (India). (2) Schonwetter 1967 (Australia).
(3) Hume and Gates 1890:268. (4) E.C.S. Baker 1929:327. (5) Medway and
Wells 1976:96.

EGG-LAYING

Place Months

N India Sep-Dec
Burma (S Pegu) Dec-Jan
Sri Lanka Early months of year
N Australia/SE Papua New Guinea Mar-Jun
SE Australia (coastal NSW) Aug-Nov

299
 Saddlebill Stork
(Ephippiorhynchus senegalensis)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 7.3 (3)

Fern. [1] — — — — — 5.0 (3)
Fern. [1] , — — — — — 6.0 (7)
Fern. [1] 675 315 375 270 — — (8)
Both [?] 600-670 273-334 311-365 250-288 up to 2700 — (1)
Both [?] 590-625 — — — — — (2)
Both [?] 630-700 — — — — (3)
—
Both [10] 600-670 273-334 311-365 260-288 — — (4)
Both [4] 620-670 282-334 293-353 257-272 — — (5)
Both [?] 650-700 320-360 330-360 270-300 — — (6)

Longevity in captivity = 19+ years (Flower 1925), 36 years (Schiiz et al. 1955).
References: (1) Brown etal. 1982:185. (2) Mackworth-Praed and Grant 1973:68. (3) F.J.Jackson 1938:76. (4)
Bannerman 1930:104. (5) McLachlan and Liversidge 1970:50. (6) Reichenow 1900-01:342. (7) O. Heinroth
1922:185. (8) Sharpe 1898:314.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
Senegal (Boghe) Jan-Feb
Extremes 12 79.0-86.3 55.3-62.0 (1) N Ivory Coast Sep-Oct
Average 12 82.5 58.5 — (1) Chad (L. Agau) Dec—Jan
Extremes 6 75.6-81.3 56.7-58.0 — (2) Ethiopia Oct-Nov
Average 6 80.0 57.0 — (2) S Sudan Sep-Apr
Calculated 6 — — 146 (2) N Uganda Nov-Jan
Extremes 9 75.6-81.3 56.0-58.0 — (3) W Uganda May-Jul
Average 9 79.1 56.7 — (3) Kenya, Tanzania Dec-Jun, Sep-Oct (mainly Mar-Apr)
Extremes — 76.0-80.5 56.0-57.5 — (4) Zambia Jan-Aug, Nov-Dec (mainly Mar-Apr)
Average — 77.0 57.0 — (5) Zimbabwe Jan-Apr, Jul (mainly Feb-Mar)
Single 1 78.0 57.0 — (6) Malawi Jan
Single 1 78.5 56.2 — (7) N South Africa (Zululand, E Transvaal) Mar-Jul
Single 1 77.3 56.0 — (7)
Single 1 76.1 56.2 — (7)
References: (1) Pitman 1965. (2) Schonwetter 1967. (3) Maclean 1985:69.
(4) Chapin 1932a:463. (5) Bouet 1955a. (6) Mackworth-Praed and Grant
1973:69. (7) McLachlan and Liversidge 1970:50.
Two eggs (in the British Museum collection), which actually belong to the
Secretarybird (Sagittarius serpentarius), are within the size-range of the Saddlebill
and were formerly misidentified as such; they may have been included in some
of the above measurements (Pitman, 1965).

300
 Jabiru Stork
(Jabiru mycterid)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [10] 625-715 295-343 285-330 (1)

Male, average [10] 660 324 308 — — — (1)
Male [4] 686-730 318-345 — — — 6.3-8.1 (2)
Male, average [4] 707 329 — — — 7.03 (2)
Male [?] 620-686 280-354 292-338 205-245 - — (8)
Male, average [?] 647 315 314 236 — (8)
Male [1] - 310 310 — — 6.7 (3)
Male [1] - 300 320 — — 7.7 (3)
Male [1] - 330 320 — — — (3)
Fern. [8] 585-665 284-328 — — — — (1)
Fern., average [8] 620 299 — — — — (1)
Fern. [2] - 267-292 — — — 4.3-5.9 (2)
Fern., average [2] 648 280 — — — 5.07 (2)
Fern. [?] 610-622 248-320 286-305 198-241 - — (8)
Fern, average [?] 614 295 297 213 - — (8)
Fern. [1] - — — - 2325 — (8)
Fern. [1] - 305 295 — — — (3)
p [1] 650 350 385 250 - — (4)
p [1] 630 — — — — — (5)
p [1] - — — — — 5.47 (6)
p 300 — - 2350 — (7)
[1] -

Longevity in captivity = 16 years (Flower 1925).

Rectal temperature = 39.1-40.5°C (Benedict and Fox 1927).
References: (1) Blake 1977:189. (2) Shannon 1987. (3) B.T. Thomas 1985:923. (4) Zotta and Fonseca 1937.
(5) Chubb in Haverschmidt 1968:28. (6) Benedict and Fox 1927. (7) Bent 1926:71. (8) Oberholser and Kincaid
1974:123-124.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
SE Mexico, Belize Dec-Jan
Extremes 8 71.5-75.3 55.0-60.5 (1) Venezuela (Llanos) Aug-Nov
Average 8 73.4 58.2 — (1) Colombia (Rio San Jorge) Oct-Nov
Average — 73.2 58.2 — (2) Guyana, Surinam Aug-Oct
E Brazil (Isla Mexiana) Jul-Aug
References: (1) Hagmann 1906. (2) Oberholser and Kincaid 1974:123. SW Brazil (Mato Grosso) Sep-Nov
The egg-measurements given in F.P. Penard and A.P. Penard (1908), and NE Argentina (Corrientes, Chaco) Aug-Oct
repeated in Schonwetter (1967), Haverschmidt (1968:28) and Kahl (1971e),
are much too large and are probably an error, referring instead to the flamingo The breeding dates given (May/June) for Surinam by
(Phoenicopterus ruber ruber) (Spaans 1975c). F.P. Penard and A.P. Penard (1908) and repeated by
Haverschmidt (1955b:32, 1968:28) and Kahl (1971e), are
probably an error (F. Haverschmidt pers. comm. 27 Jun
71, A.L. Spaans pers. comm. 5 Jul 1971, Spaans 1976b).
301
 Lesser Adjutant Stork
(Leptoptilosjavanicus)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 4.09 (1)

Male [1] — — — — — 5.23 (1)
Male [1] 575 258 225 233 — — (5)
Fern., average [?] — — — — 4.5 (2)
—
Fern. [11 263 245 258 — — (5)
—
Both [?] 580-660 260-305 228-268 230-253 — — (3)
Both, average [?] — — — — — 5.71 (4)

Longevity in captivity = 30+ years (K. Brouwer et al. pers. comm. 1990).
Rectal temperature = 38.8-40.2°C (Benedict and Fox 1927).
References: (1) J.H. Riley 1938. (2) O. Heinroth 1922. (3) E.C.S. Baker 1929:330. (4) Benedict and Fox 1927.
(5)Sharpe 1898:318.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
NE India, Burma Jul-Jan (mainly Jul-Aug)
Extremes 50 58.8-86.2 49.0-62.0 (i) S India, Sri Lanka Feb-May
Average
Extremes
50 76.4
70.5-71.5
55.3
51.8-52.8
—
—
(i) Sri Lanka
Malaysia (Johore)
Sep (sometimes)
Feb-Apr
2 (2)
Average — 73 53.5 — (3) Malaysia (Selangor) Mar-Jun
Average 3 73 55 — W Malay Peninsula Jan-Apr
Extremes 7 61.9-78.0 49.9-53.0 — (5) Sumatra (Belitung) May
Average 7 70.23 51.23 — (5) EJava Feb-Mar
Extremes 9 70.6-80.3 52.2-57.3 — (6) WJava May-Aug
Average 9 75 54.2 — (6) Samarahan Delta, Borneo May
Average — — — 113 (7)
Extremes 66 60.0-86.0 50.2-58.8 — (8)
Average 66 75.7 54.8 127 [calc.] (8)
Extremes — — 120-130 (9)
—
Average — 72.3 53.1 — (9)
References: (1) E.C.S. Baker 1935:449. (2) Hume and Gates 1890:265. (3)
Henry 1971. (4) Hoogerwerf and Rengers Hora Siccama 1937. (5) Hoogerwerf
1949b: 18-19. (6) Hellebrekers and Hoogerwerf 1967a. (7) O. Heinroth 1922.
(8) Schonwetter 1967. (9) P. Saikia (pers. comm.).

302
 Greater Adjutant Stork
(Leptoptilos dubius]

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Both [?] 800-820 320-345 320-330 310-335 - (1)

p 800 300 325 313 —
[1] (2)

Longevity in captivity = 43+ years (K. Brouwer et al. pers. comm. 1990).
References: (1) E.C.S. Baker 1929:328. (2) Sharpe 1898:317.

EGG M E A S U R E M E N T S

Length Width Weight

No. (mm) (mm) (g) Reference

Extremes 50 70.1-82.8 51.5-64.7 (i)

Average
Extremes
50
—
77.3
69-81.5
57.5
50.5-63.8
—
—
(i)
(2)
Extremes 60 71-83 53-64.7 — (3)
Average 60 78.5 57.5 — (3)
Calculated 60 — — 142 (3)
Average — 78.7 59.0 160 (4)
References: (1) E.C.S. Baker 1929:329. (2) Hume and Gates 1890:264. (3)
Schonwetter 1967. (4) P. Saikia (pers. comm.).

EGG-LAYING

Place Months

Assam, India/Burma Sep-Jan

303
 Marabou Stork
(Leptoptilos crumeniferus]

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference
a
Male [?] 705-794 279-346 266-319 278-358 2870 (1) Extremes 22 71-84.5 50-62 (1)
(max.) Average 22 79.1 55.6 — (1)
Male [?] — — 266-319 — — — (2) Calculated — — — 138 (1)
Male [37] _ — — — — 5.6-8.9 (3) Average — — — 140 (2)
Male [2] 750-765 — 290-320 — — 5.2-6.0 (5) Extremes — 80-84.5 54-57.5 140-145 (3)
Male, average PI 745 — — 323 2630 7.1 (1) Extremes — 77-84.5 54-60 _ (4)
Male, average [?] — — 293 — — — (2) Extremes 18 72.5-86.2 52.3-58.5 — (5)
Male, average [26] — — — — — 7.0 (4) Average 18 79.2 56.3 — (5)
Male, average [8] — — — — — 7.3 (4) Extremes 40 71-86.2 50-62 — (6)
Male, average [37] 745 311 293 323 — 7.06 (3) Average 40 79.1 55.9 c. 138 (6)
Male, average [9] — — — — 2630 — (3) Extremes 18 72.5-86.2 52.3-58.5 — (7)
Male [1] — — — — — 7.22 (11) Average 18 79.2 56.3 c. 138 (7)
Fern. [?] 631-709 241-314 — 240-302 — 4.0-6.8 (1) Extremes 2 81.5-84 54-57.5 — (8)
Fern. [?] — — 245-283 — — (2) Extremes 3 71-76.5 50-53.5 — (8)
— _
Fern. [22] — — — — — 4.0-6.8 (3) Average 6 80 55 (9)
Fern. [2] 620 — 205 — — 4.3-4.9 (5) Single 1 73 61 — (10)
Fern., average [?] 678 — — 282 2470 5.7 (1) Extremes 14 73-84 52-62 — (11)
Fern., average [?] — — 266 — — — (2) Average 14 79.5 56.0 — (11)
Fern., average [14] — — — — — 5.7 (4) Calculated 14 — — 138 (11)
Fern., average [6] — — — — — 5.8 (4)
Fern., average [22] 679 278 266 282 — 5.66 (3) References: (1) Brown et al. 1982:190. (2) O. Heinroth 1922. (3) Reichenow
Fern., average [7] — — — — 2470 — (3) 1900-01:341. (4) Chapin 1932a:468. (5) W. Fraser 1971. (6) Maclean 1985:70.
Fern. [1] — — — — — 6.12 (5) (7) MacLachlan and Liversidge 1970:47. (8) M.E.W. North 1943. (9) Pomeroy
Fern. [1] — — — — — 6.7 (11) 1977b. (10) FJ. Jackson 1938:80. (11) Schonwetter 1967.
Fern. [1] — — — — — 4.82 (11)
Both [?] 643-656 225-281 225-248 244-256 — — (6)
Both [?] 650-700 — — — — 7.4 (7)
Both [10] 620-720 226-280 215-270 250-295 — — (8) EGG-LAYING
Both [?] 650-700 240-280 250-270 250-300 2540 — (9)
Both, average [10] 673 244 242 276 — — (8)
P [1] 725 270 275 295 — — (10) Place Months
a
Reputed spans of 4 m or over can be discounted (Brown et al. 1982:187). N South Africa (Transvaal) May
Longevity in captivity = 41+ years (K. Brouwer et al. pers. comm. 1990). NE Swaziland Jul
Resting body temperature = 39.2-41°C (Bamford and Maloiy 1980). Zimbabwe May-Sep
Mean resting heart-rate = 90.6 ± 1.8 beats/min (Bamford and Maloiy 1980). N Botswana Jul-Sep
References: (1) Brown et al. 1982:187. (2) Maclean 1985:69. (3) Pomeroy 1977b. (4) Pomeroy 1973. (5) K.M. S Tanzania, N Zambia, Malawi May-Sep
Schneider 1952. (6) Heuglin 1869-73:1115. (7) F.J.Jackson 1938:77. (8) MacLachlan and Liversidge 1970:46. N Tanzania, SE Kenya Jul-Sep
(9) Reichenow 1900-01:338. (10) Sharpe 1898:320. (11) FitzPatrick Inst., Univ. of Cape Town (unpubl. data). NE, W Kenya Sep-Jan
Uganda, E Zaire Oct-Jan
S Sudan Sep-Jan
S Ethiopia, Somalia Oct-Nov
N Nigeria Aug-Mar
S Chad Nov-Jan
Gambia Dec-Jan

304
 Shoebill
(Balaeniceps rex)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [1] 780 191 245 258 (7)

Male [1] 705 213 246 281 — — (8)
Male [1] 710 238 255 284 — 6.70 (4)
Fern. [3] — — — — 5.4-5.9 (1)
—
Fern., average [3] — — — — — 5.57 (1)
Fern. [1] 655 208 217 276 — 4.36 (4)
Fern. [1] 667 — _ — — (8)
—
Fern. [1] 700 — — — 2605 — (9)
Both [?] 588-700 188-200 221-244 244-260 — — (2)
Both [?] 655-685 — — — — — (3)
Both [4] 660-700 228-235 254-255 279-305 — —
__ (4)
Both [?] 655-780 — — — — (5)
Both, average [?] 707 — — — — — (5)
Both, average [?] 700 230 250 280 — — (6)
? [1] 650 203 250 250 — — (10)
p 721 — — — — — (8)
[1]
p 730 245 — 250 2620 — (4)
[1]
Longevity in captivity = 36 years (Flower 1938:220, Bourliere 1946).
References: (1) H.G. Klos 1985. (2) Heuglin 1869-73:1096. (3) Mackworth-Praed and Grant 1973:62. (4) W.
Fischer 1970:13. (5) Maclean 1985:63. (6) Reichenow 1900-01:357. (7) Bannerman 1930:87. (8) Chapin
1932a:449. (9) FJ. Jackson 1938:63. (10) Sharpe 1898:287.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
S Sudan Sep-Feb (mostly Nov-Jan)
Extremes 10 80.1-90.0 56.9-61.3 (1) Uganda (L. Kyoga, Rwenzama Swamp) Mar-Jun
Average 10 85.7 59.4 — (1) SE Zaire (L. Kabamba) Aug-Sep
Calculated 10 — — 164 (1) NE Zambia (Bangweulu) Apr-Jul
Extremes 3 81.0-82.0 60.0-61.0 — (2)

References: (1) Schonwetter 1967. (2) F.J.Jackson 1938:66.

305
 American White Ibis
(Eudocimus ruber)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [6] 266-289 156-169 94.97 (1)

Fern. [6] 251-269 123-131 — 84-95 — — (1)
Male [5] 279-285 145.4-163.0 90.3-102.5 107.1-119.8 — — (1)
Fern. [5] 260-268 118.5-130.0 83.8-88.0 93.3-104.8 — — (1)
Male, average [6] 282.3 163.2 — 95.6 — — (1)
(ruber}
Fern., average [6] 260.6 127.9 — 90.2 — — (1)
Male, average [5] 282.6 155.1 95.9 121.1 — — (1)
(albus)
Fern., average [5] 265 125.6 85.5 98.6 — — (1)
? (ruber) — 254 110 — 114 — — (2)
? (albus) — 279 162 89 114 — — (3)
Male [7] 205-295 136-164 — — —
_ —
_ (4)
Male, average — 277.3 153.3 — — (4)
Male [1] 310 153 — — — — (4)
Fern. [9] 263-278 119-130 — — - 0.772-0.797 (4)
Fern., average P] 271.5 124 — — — — (4)
Male [2] 258-263 157-168 — — - 0.935 (5)
Fern. [2] 258-263 120 — — — — (5)

References: (1) Blake 1977:199. (2) D.G. Elliot 1877. (3) D.G. Elliot 1877. (4) Palmer 1962:524. (5) Palmer
1962:530.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
S Florida Mar-Apr, or later up to Sep
20 57.53 38.65 (1) Carolinas Apr-May or later
Venezuela (Llanos) Jun/Jul onwards
Reference: (1) Palmer 1962:528. Trinidad Apr-Sep
Surinam Apr-Aug
French Guiana Apr-Jun
Brazil Mar-Sep

Nesting is highly variable, depending on water con-

ditions; on the South American coast it is mostly during
the wet season. Elsewhere nesting is either during rising or
falling water levels.

306
 Barefaced Ibis
(Phimosus infuscatus)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

P. i. berlepschi
Male [8] 270-292 114-127 - 122-129 - - (1)
Male, average [8] 281.3 120.2 - 125.7 - - (1)
Fern. [8] 252-275 106-121 - 110-124 - - (1)
Fern. , average [8] 265.2 108.8 - 114.1 - - (1)

P. i. nudifrons
Male [10] 273-299 104-136 - H9_134 _ _ (1)
Male, average [9] 285 127.5 - 124.2 - - (1)
Fern. [8] 251-295 101-112 - 105-130 (1)
Fern., average [8] 264.6 108.9 117.7 - - (1)

P. i. infuscatus
Male [10] 270-289 104-124 - 116-131 (1)
Male, average [10] 282.1 113.6 121.7 - - (1)

Reference: (1) Blake 1977:198.

EGG MEASUREMENTS

Length Width Weight

No. (mm) (mm) (g) Reference

P. i. nudifrons 45.0 33 28 (1)

P. L berlepschi 48.4 32 27.5 (1)

References: (1) Schonwetter 1967.

EGG-LAYING

Place Months

Venezuela
(Llanos) May-Oct
Brazil Oct-Nov

Nesting is prolonged and

continuous throughout the
rainy season.

307
 Glossy Ibis
(Plegadis falcinellus]

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

Male [8] 275-294 127-129 99.2-112.0 98-108 (1)

(1)
133
13
45-59
47.5-57.6
33.40
31.5-38.2
38
—
(i)
Male, average [8] 288.3 131 103.1 102.3 (2)
Fern. [10] 252-275 98-114 78-88 90-100 (1) — 49.3-56.1 33.8-37.6 — (2)
Fern., average [10] 262.2 104.2 81,8 93.6 (1) 100 52.18 36.9 — (3)
— 54 35 — (4)
INDIA 13 48.9-54.7 35.5-37.3 — (5)
Both — 248-298 99-144 85-100 94-106 (2)
References: (1) Brown etal. 1982:194. (2) Marchant and Higgins 1990:1075.
EUROPE _ (3) Baker in S. Ali and S.D. Ripley 1968:115. (4) Mackworth-Praed and Grant
Male — 290-306 — — - 0.750 (2) 1973: Series 1, Vol. 78. (5) McLachlan and Liversidge 1970:P56.
Fern. — 264-277 — — — (2)

AUSTRALIA
Male — 275-294 126.3-140.1 96.8-107.7 93-107 (3) EGG-LAYING
Fern. [5] 265-275 97.8-112.7 67.5-86.3 96-106 - 0.485-0.970 (3)
Fern., average [5] 267 106.3 79.4 98.8 (3)
Fern., average [5] 259.8 — — — (4) Place Months
Male, average [5] 286 — — — (4)
Male [7] 280-306 126-141 101-113 96-111 (5) India May-Jul (but sometimes in rains) Jan—Feb
Southeast Asia, Burma (formerly) May—Jun
EUROPE Australia (NSW Balvanald)a Oct-Nov
Male, average [7] 297 132 107 106 (5) Australia (Cowal)a Oct-Feb
Fern. [7] 267-281 106-114 82-90 90-99 (5) Algeria, Morocco, Egypt Jun (previously)
Fern., average [7] 273 110 86 94 (5) Mali Mar, Jul-Sep
? 257-304 114-136 73-111 — (6)
— Kenya May-Aug, Jan-Mar (short rains)
Fern. [?] 250-275 100-135 — — (7) Tanzania Apr
Zambia Feb
References: (1) Blake 1977:201. (2) S. Ali and S.D. Ripley 1968:115. (3) Marchant and Higgins 1990:1077. (4) Namibia Aug
Amadonand Woolfenden 1952 in Marchant and Higgins 1990:1077. (5) Cramp 1977:343. (6) Palmer 1962:517. Zimbabwe Jan
(7) Witherby et al. 1939 in Palmer 1962:517. South Africa (SW Cape) Sep-Nov
South Africa (Transvaal) Sep-Jan, Oct-Dec
NEUSA May-Aug
Florida, USA Mar-Jun
West Indies Feb-May
Venezuela Feb-May
Trinidad & Tobago
Europe May-Aug

Nesting highly variable and sporadic throughout the extensive range.

a
Nest times depend on rainfall and nesting periods of other ibises, egrets and spoonbills.

308
 Whitefaced Ibis Puna Ibis
(Plegadis chihi] (Plegadis ridgwqyi)

BODY M E A S U R E M E N T S BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Referei

Male [11] 261-280 118-143 91-98 92-100 - - (1) Male [10] 292-302 108-127 80-88 117-125 - - (1)
Male, average [11] 269.6 133.9 94.6 93.7 - - (1) Male, average [10] 298.4 117.4 84 120.2 - - (1)
Fern. [8] 238-249 100-111 63-73 83-94 - - (1) Fern. [7] 266-287 86.95 59-68 106-112 - - (1)
Fern., average [8] 243.5 103.6 70 87.8 - - (1) Fern., average - 276.8 92 63.5 109 - - (1)
Reference: (1) Blake 1977:200 Reference: (1) Blake 1977:202.

EGG M E A S U R E M E N T S EGG MEASUREMENTS

Length Width Weight Length Width Weight

No. (mm) (mm) (g) Reference No. (mm) (mm) (g) Reference

Average - 50.8 35.8 36 (1) Average - 52 35.4 35 (1)

Extremes - 42-55 32.4-39.4 - (1) Extremes - 49.0-55.4 33.2-37.3 - (1)

Reference: (1) Schonwetter 1967. Reference: (1) Schonwetter 1967.

EGG-LAYING EGG-LAYING

Place Months Place Months

USA Apr-May, Jun (occasionally) Peru Apr-Jul, Jan-Feb

South America Nov-Dec Ayasuchu Jan-Mar
South Bolivia Nov-Jan/Feb

309
 Sharptailed Ibis Plumbeous Ibis
(Cercibis oxycercd) (Harpiprion caerulescens]

BODY MEASUREMENTS BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) "(kg) Reference

Male [9] 390-415 156-168 - 250-301 (1) Male [10] 397-450 145-167 - 165-190 - - (1)
Male, average [9] 399.7 161.6 - 272.6 (1) Male, average [10] 413.4 153.8 — 179.5 - — (1)
Fern. [4] 376-413 144-167 - 256-272 (1) Fern. [10] 360-406 126-147 - 165-192 - - (1)
Fern., average [4] 394.2 152.7 - 262.2 (1) Fern., average [10] 384.8 134.6 - 176 - - (1)
Reference: (1) Blake 1977:195. Reference: (1) Blake 1977:192.

EGG M E A S U R E M E N T S EGG M E A S U R E M E N T S

Length Width Weight Length Width Weight

No. (mm) (mm) (g) Reference No. (mm) (mm) (g) Reference

Average — 65.9 44 70 (1) Average - 68.6 44.9 74 (1)

Extremes
1? t-*~a.*-^t* — o4.
C^A, 1/—OD.J
££ t% 4-z./—
4 O 7 44. o
A.A. Q
(1)
Reference: (1) Schonwetter 1967.
Reference: (1) Schonwetter 1967.

EGG-LAYING
EGG-LAYING

Place Months
Place Months
Mato Grosso Mar/Apr-Oct
Llanos (S America) Aug/Sep-Feb
Corresponding with commencement of
Nesting is in the dry season. the dry season.

310
 Buffnecked Ibis
(Theristicus caudatus]

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Caudatus
Male [10] 385-429 131-160 - 193-220 - - (1)
Male, average [10] 402.7 149 - 204.8 - - (1)
Fern. [10] 378-405 131-150 - 180-203 - - (1)
Fern., average [10] 389.8 138.8 - 190.9 - - (1)
Hyperorius
Male [7] 395-418 143-172 - 196-224 - - (1)
Male, average [7] 406.3 156.8 - 209.3 - - (1)
Fern. [6] 384-410 140-156 - 194-211 - - (1)
Fern., average [6] 395.1 146.8 - 202.8 - - (1)
Branickii
Male [7] 391-407 113-134 - 187-203 - - (1)
Male, average [7] 400.7 126.5 - 195.5 - - (1)
Fern. [6] 375-399 109-130 - 178-199 - - (1)
Fern., average [6] 388.8 120 - 186 - - (1)
Melanopis
Male [6] 374-398 120-151 - 169-191 - - (1)
Male, average [6] 383 138.5 - 179.8 - - (1)
Fern. [7] 351-381 113-132 - 158-180 - - (1)
Fern., average [7] 368.4 122.6 - 172.3 - - (1)
Reference: (1) Blake 1977:194.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
South Chile Sep-Dec
Average -
-
66.9
63.6-68.6
44.3
44.0-44.6
69 (i) Argentina Nov-Feb
(2) Tierra del Fuego Nov-Feb
Peru Jun
References: (1) Schonwetter 1967. (2) Kreugerin Schonwetter 1967. Llanos (Paraguay, Brazil, Venezuela) Mar-Oct (but found all the year round depending on climate
and altitude)

311
 Green Ibis
(Mesembrinibis cayennensis)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [51 263-300 104.5-118.5 61.8-63.5 127-155 (1)

Male, average [51 280 112.6 62.4 138 — — (1)
Fern. [41 280-289 103-113 56.8-62.2 131-148 — — (1)
283 108.5 59.5 142 — — (1)

Reference: (1) Blake 1977:196.

EGG M E A S U R E M E N T S

Length Width Weight

No. (mm) (mm) (g) Reference

Average 54.6 38.6 44 (i)

Extremes - 53-57 37.1-39.5 (i)
Reference: (1) Schonwetter 1967.

EGG-LAYING

Place Months

Panama Mar-Apr
Colombia Mar-Apr
Surinam Sep
Venezuela Jun-Aug

312
 Hadada Ibis
(Bostrychia hagedash}

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

B. h. hagedash
Both [17] 334^370 117-153 63-73 137-170 - 1.262 (1)
Average - 353 134 68 154 (1)
B. h. brevirostris
Both - 330-383 126-163 (1)
B. h. nilotica
Both - 335-392 152-174 - - - - (1)
sexes alike

B. h. brevirostris
Both [5] 335-365 130-141 67-73 145-153 - - (2)
References: (1) Brown et al. 1982:195. (2) Gates 1930:117.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
Guinea-Bissau May-Oct (rains)
172
—
54-66 38-48
42
55-62
—
(i) Gambia Jan-Mar (dry)
Jun
62 (2) Niger
— 51 45 — (2) Nigeria Jun-Jul (rains)
90 57.0-65.1 38.5-46.8 — (3) Ethiopia Sep
Kenya, Uganda Mar-Jun (peak long rains; but can be year round)
References: (1) Brown et al. 1982:196. (2) Mackworth-Praed and Grant Tanzania Apr-Jul, Oct
1973:75. (3) McLachlan and Liversidge 1970:56. Zambia Jan, Mar-Apr, Jul-Dec
Zaire Jan-Dec
Zimbabwe Oct-Dec
Mozambique Dec
South Africa Jan-Dec

This species has an extended breeding season throughout the year but peak
laying occurs during and after the main rains.

313
 Wattled Ibis
(Bostrychia carcunculatd)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Both [7] 358-380 110-127 63-66 150-180 - - (1,2)

References: (1) Moltoni and Ruscone 1942. (2) Friedman 1930 in Brown et al. 1982:196.

EGG M E A S U R E M E N T S

Length Width Weight

No. (mm) (mm) (g) Reference

- 59-61 40-59 50 (1)

Reference: (1) Brown et al. 1982:197.

EGG-LAYING

Place Months

Ethiopia Mar-May, Jul, Dec

314
 Olive Ibis
(Bostrychia olivacea)

BODY M E A S U R E M E N T S EGG M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

B. o. olivacea B. o. akeleyorum — 56-58 40-41 50 (1)

Male — 333 96 67 146 — — (1) (Type) Guinea Coast B. o. akeleyorum — 57.1 40.2 50 (2)
Male — 330 c. 100 72 150 — — (1) (Type of splendida) Liberia
Ad. — 330 95 73 150 — — (2) Bannerman collection No data available on other races.
Ad. — 334 96 68 147 — — (2) Bannerman collection References: (1) Brown et al. 1982:198. (2) Schonwetter 1967.

B. o. akeleyorum
Male — 358 108 71 160 - - ( 1 ) (Type) Mt. Kenya 9000 ft
(Akeley) EGG-LAYING
Fem. — 362 102 64 165 - - ( 1 ) Mt. Kenya 9000 ft (Akeley)
Male — 355 115.5 73 150 - - ( 1 ) Mt. Kenya 6500 ft (van
Someren) Place Months
B. o. rothschildi Kenya Jun-Aug
Male - 328 95 66 (1) (Type) Prince's Is. (L. Fea)
Fem. — 313 c. 90 67 110 (1) Prince's Is. (Dohrn and
This species is little known,
Keulemans)
with no breeding records outside
B. o. cupreipennis
Kenya.
Male (?) - 310 85 57 130 (1) (Type) Bipindi (Zenker)
Imm. — 280 76 (1) Efulen (Bates)
Ad. 317 86.5 62 125 (1) Gamma R. (Du Chaithu)
Ad. - 309 85 59 127 (1) Near Bolobo (Schouteden)
Fem. - 335 94 70 133 (1) Avakubi (Lang and Chapin)

B. o. bocagei
Ad. - 255 75 53 103 - - (1) (Type) Sao Tome
Male - 256 76 55 101 — ( 1 ) Sao Tome (F. Newton)
Male - 233 75 50 88 - ( 1 ) Sao Tome (F. Newton)
Fem. - 255 68 50 96 - - ( 1 ) Sao Tome (F. Newton)
p - 250 75 — — — (3)

References: (1) Chapin 1923. (2) de Naurois 1973. (3) Specimen in British Museum of Natural History.
Measurements taken by Chapin were with the wing straightened and this is probably the case in other reports,
but slight differences occur, with some authors using the same skins. There are now estimated to be 9 skins only of
B. o. bocagei:, in spite of this small number it is to be hoped that should this race be re-discovered the temptation to
collect further specimens will be resisted.

315
 Spotbreasted Ibis Madagascar Crested Ibis
(Bostrychia rara) (Lophotibis cristatd)

BODY M E A S U R E M E N T S BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex W (mm) (mm) (mm) (mm) (mm) •(kg) Reference

Both [10] 270-290 115-130 56-65 112-120 — (1) Both - 355 134 64 184 - - (1)
Male [3] 265-291 122-131 57-63 102-120 - (2)
Fern. [3] 253-275 122-131 57-63 102-120 (2) Reference: (1) Elliott 1877:500.

References: (1) Brown et al. 1982:198. (2) Gates 1930:118.

EGG M E A S U R E M E N T S

EGG M E A S U R E M E N T S
Length Width Weight
No. (mm) (mm) (g) Reference
Length Width Weight
No. (mm) (mm) (g) Reference 55 41 (1)

Average 55 35 (1) Reference: (1) Langrand 1990:117.

Reference: (1) Brosset and Erard 1976.

EGG-LAYING

EGG-LAYING
Place Months

Place Months Madagascar Sep-Jan

Gabon Jan-Jun, Sep-Dec

Throughout West Africa Mar-Jun, Sep-Dec
Nest building carried out when flood waters are high.
Laying coincides with rainfall peaks, but some may occur
if water remains high in dry season.

316
 Sacred Ibis
(Threskiornis aethiopicus)

BODY MEASUREMENTS EGG-LAYING

Wing Culmen Tarsus Tail Wing-Span Weight Place Months

Sex M (mm) (mm) (mm) (mm) (mm) (kg) Reference
Aldabra (Seychelles) Nov-Dec
Male [2] 380-385 162-183 100-116 129-140 1120-1240 1.53 (1,2) Senegal Nov-Dec
Male, average [2-5] 383 170 108 134 — — (1,2) Gambia Jan, Jul, Nov-Dec
Fern. [3] 361-362 135-157 94-103 129-135 — — • (1,2) Guinea-Bissau Mar-Sep
Fern., average [3-81 362 146 98 131 — — (1,2) Nigeria Jan-Feb, May-Jun, Dec
Male [1] 396 167 105 140 — — (3) Mali Jul-Feb
Chad Jun
References: (1) Brown et al. 1982:200. (2) Cramp 1977:351. (3) Gates 1930:118. Sudan Mar, Jun-Aug
Ethiopia Mar-Jun, Aug-Sep, Jan-Jun
W Kenya Apr
Central Kenya May-Jun, Aug, Nov
EGG MEASUREMENTS Uganda Feb, May, Nov
Burundi Jul
Tanzania Jan-Feb, May, Oct
Length Width Weight Zambia Jan-Sep
No. (mm) (mm) Reference Malawi Jan-Sep
(g)
Zimbabwe Apr
T. a. aethiopicus _ 56-70 38-48 62 Namibia Jul-Aug
(1)
42 58-68 39-46 62 (2) South Africa Jan-Mar, May, Aug-Dec
— 64 42 — (3)
60 60.2-70.5 39.7-51.2 — (4)

T. a. bernieri
Average — 59.5 41.3 55 (5)

References: (1) Brown et al. 1982:201. (2) Schonwetter 1967 in Cramp

1977:350. (3) Mackworth-Praed and Grant 1973:73. (4) McLachlan and
Liversidge 1970:54. (5) Schonwetter 1967.

317
 Oriental White Ibis Australian White Ibis
( Threskiornis melanocephalus] ( Threskiornis moluccd)

BODY M E A S U R E M E N T S BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Both - 343-370 139-170 99-118 133-145 - - (1) Male [3] 355-398 183.5-197.1 94.9-108.1 120-134 1100-1250 1.7-2.5 (1)
Male - 381 185 119 129 - - (1) Male, average [3] 380.6 188.6 102.7 126.3 - - (1)
rem.
T?A<~% [4J
r/ll "2Ei£ Q"7O 149.1—
5D3—672 1/iO 1 15o.4
1 £Q A 9^.4—95.4
QO A O£ A \2.i—
1 OO 150
1 QA — — (1)
Reference: (1) S. Ali and S.D. Ripley 1968:1 12. Fern., average [4] 363.7 153.7 94.1 125 - 1.4-1.9 (1)
Male - 380-397 186-190 94-107 - - - (2)
Male, average - 389 188 103 - - - (2)
Fern. - 348-373 131-157 81-90 - - - (2)
EGG MEASUREMENTS Fern., average — 361 148 85.8 — — — (2)
Male - - 173-210 - - - - (3)
Male average 190 5 (3)
Length Width Weight Fern. - - 140-168 - - - - (3)
No. (mm) (mm) (g) Reference Fern., average — — 154 — — — — (3)

150 63.5 43.1 - (1) References: (1), (2), (3) in Marchant and Higgins 1990:1086. (1) SAM, MV. (2) Healesville Fauna Reserve
Average 11 62.31 41.43 - (2) Victoria (ABBBS). (2) Vic. (K.W. Lowe).
11 59.4-65.1 39.2-44.2 5.073 (2)

References: (1) Baker in S. Ali and S.D. Ripley 1968:111. (2) Hellebrekers
and Hoogerwerf 1967: 14. EGG MEASUREMENTS

Length Width Weight

EGG-LAYING No. (mm) (mm) (g) Reference

60 71 41-47 668 (1)

Place Months - 59-75 39-49 - (1)
59 78 39-49 (1)
N India Jun/Jul-Oct - 60-70 40-46 - (1)
S India, Sri Lanka Nov-Feb/Mar
References: (1) Marchant and Higgins 1990:1084.

EGG-LAYING

Place Months

S Victoria (Australia) Jun-Nov, May-Mar (rarely)

S Australia Sep-Dec, Jan
NSW (Australia) Sep-Apr

This species does not always breed annually and variations of

nesting times occur according to rainy season.

318
 Strawnecked Ibis Waldrapp Ibis
( Threskiornis spinicollis] (Geronticus eremita)

BODY MEASUREMENTS E&ODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male - 364-415 158.2-179.8 83.6-102 130-152 - 1.4-1.57 (1) Male - 403-420 133-147 68-72 196-220 - - (1)
Fern. - 351-375 133.4-142.2 78.4-83.2 126-142 - 1.15-1.32 (1) Fern. - 390-408 115-131 68-72 196-220 - - (1)
Reference: (1) Marchant and Higgins 1990. Western population (Morocco)
Male - - 132-146 (1)
Fern. - - 128-145 - - - - (1)

EGG MEASUREMENTS Northeastern population (Turkey)

Male - - 126-132 (1)
Fern 121 127 (\\
\l)
Length Width Weight
No. (mm) (mm) (g) Reference Reference: (1) Brown et al. 1982.

14 61.4-69.1 42.4-47.0 - (1)

Average - 64.8 44 69 (2)
Extremes - 57.5-70.0 38.6-48.0 - (2) EGG M E A S U R E M E N T S
f
References.
"D f^»-^v-.^^r- • (1)
/ 1 \ Marchant
A/f ^*~^\->^n + and rliggms
~nA tTi/- lyyu.
r /- r 4v-,r- 1 QQfV 1lUyo.
flO*-? ((t) ocnonwetter 1Q£7
)\ Qr«J-»orMA7*»t*-«»»- iyt>/.
Length Width Weight
No. (mm) (mm) (g) Reference

EGG-LAYING 46 61-69 42-46 68 (1,2)

13 - - 54-67 (3)
63 44 (4)
Place Months
References- (1) Brown et al 1982*204 (2) Schonwetter 1967 in Cramp
S, SE Australia Sep-Feb 1977:347. (3) Wackernagel 1964 in Cramp 1977:347. (4) Mackworth-Praed and
W Australia Sep-Feb Grant 1973:74.
N Australia Sporadic not annually
This species is largely dependent on rainfalls but
uses permanent water also. It often nests in com-
EGG-LAYING
pany with Australian White Ibis.

Place Months

Morocco Mar-May
Algeria Only after successful rains

319
 Bald Ibis Black Ibis
( Geronticus calvus) (Pseudibis papillosa)

BODY M E A S U R E M E N T S BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) • (mm) (mm) (mm) (kg) Reference

Both [7] 369-403 140-152 66-73 175-200 - - (1) P. p. papillosa

Both — 365-400 138-158 75-85 165-194 — — (1)
Note: females are slightly larger with a slightly longer bill and tarsus.
Reference: (1) Brown et al. 1982. P. p. davisoni
Both - 419 197 83 229 - - (2)

References: (1) Baker (in Ali and Ripley 1968). (2) D.G. Elliott 1877:49.
EGG MEASUREMENTS

Length Width Weight EGG M E A S U R E M E N T S

No. (mm) (mm) (g) Reference

51-71 38-49 68 (1) Length Width Weight

24 57-71 38-49 - (2) No. (mm) (mm) (g) Reference

P. p. papillosa
References: (1) Brown et al. 1982:206. (2) McLachlan and Liversidge Average - 63.0 43.8 - (1)
1970:55. Average - 63.0 43.8 64 (2)
Extremes - 56-70 38-50 - (2)

P. p. davisoni
EGG-LAYING Average - 63.7 45.2 71 (2)
Extremes - 61.0-68.2 43.9-46.7 - (2)

Place Months References: (1) Baker in S. Ali and S.D. Ripley 1968:133. (2) Schonwetter
1967
South Africa Aug-Oct

EGG-LAYING

Place Months

P. p. papillosa
N India Mar and Oct
Gujarat, India Apr-May, Dec-Jan
S India Apr-May, Dec-Jan
Nepal Mar and Oct
In all areas two distinct nesting periods occur,
but these are extremely variable due to rains.

P. p. davisoni
Burma (formerly) Feb-Mar

320
 Giant Ibis Oriental Crested Ibis
(Thaumatibis giganted) (Nipponia nippori)

BODY M E A S U R E M E N T S BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male - 546 236 112 267 (1) Both — 419 165 89 165 (1)
Fern. 546 241 108 279 (2)
Reference: (1) D.G. Elliott 1877.
References: (1) Robinson and Kloss 1911. (2) D.G. Elliott 1877.

EGG MEASUREMENTS
EGG MEASUREMENTS

Length Width Weight

Length Width Weight No. (mm) (mm) (g) Reference
No. (mm) (mm) (g) Reference
Average 57.3 35.6 39(?) (1)
Not recorded Extremes - 50.5-66.0 31.5-40.5 (2)
References: (1) Schonwetter 1967. (2) Swinhoe and Nehrkorn in Schonwetter
1967.

EGG-LAYING
EGG-LAYING

Place Months

Not recorded Place Months

China Mar
Japan (formerly) mid-Apr

321
 Eurasian Spoonbill
(Platalea leucorodid)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight Wing Culmen Tarsus Tail Wing-Span 'Weight
Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

P. I. leucorodia P. I balsaci
Male [14] 367-409 194-222 148-150 — ' — — (1) Male [9] 364-390 157-200 123-144 - (1)
Male, average [14] 389 208 149 — — — (1) Male, average [9] 372 184 130 - (1)
Male [3] 382-401 211-224 144-153 121-126 - — (2) Male [1] 382 205 140 132 (11)
Male, average [3] 394 216 149 124 — (2) Fern. [8] 340-362 151-180 115-129 - (1)
Male [13] 386-412 195-231 140-163 108-126 - — (3) Fern., average [8] 352.5 167 120 (1)
Male, average [13] 394 213 149 117 - — (3)
Male [8] 350-388 175-223 113-150 — — — (16) P. I. archeri
Male, average [8] 373 206 141 — — — (16) Both [?] 325-360 145-170 102-118 - (3)
Male [5] 375-395 205-230 140-150 110-120 - — (12) Unsexed [1] 325 153 103 - (13)
Male [1] — — — — — 1.96 (8) Unsexed [1] 308 150 113 - (16)
•Male
\,ri 2 .DJ
/-vr /n\
U)
Male [1] — — — — — 2.08 (2) Longevity in wild = 28 years 2 months (Rydzewski 1973).
Male [1] — — — — — 1.656 (3) Longevity in captivity = 27 years (Schenker 1978).
Male [1] 394 238 158 132 - — (10) References: ( 1 ) de Naurois and Roux 1974. (2) T.H. Shaw 1936:128-129. (3) Cramp 1977:352, 357. (4) E.C.S.
Male [1] 410 230 145 — — — (13) Baker 1929:312. (5) Dementievand Gladkov 195 1:421. (6) Hartert 1912-21 :1217-1218. (7) Lippens 1954:26. (8)
Fern. [22] 355-388 176-223 133-148 — — — (1) Vasvari 1955. (9) S. Ali and S.D. Ripley 1968:117. (10) La Touche 1931-34:431. (11) de Naurois 1959. (12)
Fern., average [22] 370 191 139 — — — (1) Witherbyetal. 1939: 121.(13)Pers.obs.M.P.K. (Leiden Mus./26 Apr 89). (14) Kobayashi 1963:67. (15) Sharpe
Fern. [2] 364-369 187-194 134-139 110-113 — (2) 1898:46-47. (16) Ogilvie-Grant 1889.
Fern., average [2] 367 191 137 112 - — (2)
Fern. [10] 360-377 168-191 123-141 108-118 - — (3)
Fern., average [10] 370 182 131 113 - — (3)
Fern. [10] 350-373 168-183 115-140 — — — (16)
Fern., average [10] 359 177 129 — — — (16)
Fern. [8] 350-370 175-190 — — — — (12)
Fern. [1] — — — - 1258 — (5)
Fern. [1] — — — 1400 — (5)
Fern. [1] — — — - 1450 — (5)
Fern. [1] — — — — — 1.94 (2)
Fern. [1] 365 183 125 — — — (13)
Both [?] _ — — - 1150-1300 — (3)
Both [?] 350-395 180-228 130-165 108-122 - — (4)
Both [?] 360-397 183-223 120-145 — — — (6)
Both [?] 350-410
_ 180-238 130-165 110-132 - — (14)
Both [?] — 140-150 — — — (7)
Both [12] 370-410 — — — — — (5)
Both, average [12] 387.5 — — —
_ —
_ — (5)
Both [?] — 173-225 128-163 — (15)
Both, average [?] — 199 143 — — — (15)
Both, average [?] 373 203 143 118
_ _ — (15)
Unsexed [1] — — — 1.77 (9)
Unsexed [1] — — - — — 1.785 (9)

322
Eurasian Spoonbill continued

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
P. I. leucorodia
P. L leucorodia Spain, Portugal Feb-Apr
Extremes 250 56-74.5 40.0-49.5 — (i) N, central Europe Apr-Jun
Average 250 67.2 45.5 - (i) N India, Pakistan Jul-Oct
Calculated
Extremes
250 —
47.9-77.1
-
36.0-51.7
76 (i) S India
Sri Lanka
Nov—Jan
Dec-Apr
105 (2)
Average 105 61.2 42.4 - (2) Algeria (formerly: 19th century) May
Extremes 17 65.1-73.0 43.2-49.0 - (3)
Average 17 69.5 45.0 - (3) P. L balsaci
Extremes 40 61.1-72.1 41.0-47.7 — (4) Mauritania Apr-Jul
Average 40 65.6 44.2 - (4)
Extremes 100 62.2-78.3 41.5-49.5 - (5) P. I. archeri
Average 100 68.2 46 - (5) Red Sea, Somalia (coastal) Apr-May, Jun-Aug
Extremes 100 58.2-74.6 41-49.2 - (7)
Average 100 67.2 45.4 - (7)
Extremes 160 58.5-76.4 41.2-49.0 - (8)
Average 160 67.4 46.8 - (8)
P. L balsaci
Average 12 67 45 (9)
P. L archeri
Extremes 15 57-65 40-45 - (6)
Average 15 60 43.2 - (6)
References: (1) Schonwetter 1967. (2) Vespremeanu 1968a. (3) Dementiev
and Gladkov 1951:418. (4) E.C.S. Baker 1929:312. (5) Hellebrekers 1967. (6)
Archer and Godman 1937:82. (7) Witherbyetal. 1939:119. (8) Bauer and Glutz
von Blotzheim 1966:438. (9) Etchecopar and Hue 1967:63.

323
 Royal Spoonbill
(Platalea regid)

BODY MEASUREMENTS EGG MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight Length Width Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference No. (mm) (mm) (g) Reference

Male M 355-370 165-198 120-143 (i) Extremes 16 59.5-69.3 41.7-44.5 (1)

Male, average [4]
Male
361
353-374
187
180-196
133
120-143
—
—
—
—
—
—
(i) Average
Calculated
16
16
65.7
—
43.2
— 66
(1)
(1)
[3] _ _ (2)
Male, average [3] 362 190 134 — (2) Extremes 18 58.5-69.6 40.1-44.9 (7)
Male [3] — 183-196 121-137 — — — (4) Average 18 63.6 43.3 (7)
Male, average [3] — 188 129 — — — (4) Average — 65 44 (2)
Male [3] — 176-187 — — — — (3) Average — 64.3 42.8 (4)
Male, average [3] — 180 — — — — (3) Average 8 65.5 42.7 (5)
Male [6] 347-385 205-219 116-127 — - 1.65-2.07 (15) Average 16 68 45 (6)
Male, average [2] — 183 130 — — — (4) Extremes 11 60.5-64.5 42.8-44.5 (3)
Male [1] — — — — 1200a - (4) Ova)
Male [1] 370 168 121 — —
_ —
_ (8) Average 11 62.5 43.5 (3)
Male [1] 390 189 130 — (8) Java)
Male [1] — 188 135 — _ _ (10)
Fern. [5] 325-358 163-175 113-125 — — — (1) References: (1) Schonwetter 1967. (2) Beruldsen 1980:158. (3) Hoogerwerf
Fern., average [5] 339 166 120 — — — (1) 1952. (4) Lucas and LeSouef 1911:126. (5) AJ. North 1913-14:13. (6) Vestjens
Fem. [9] — 136-189 — — — — (3) 1977b. (7) Hellebrekers and Hoogerwerf 1967:14.
Fern., average [9] — 164 — — — — (3)
Fem. M — 164-172 115-128 — — — (9)
Fem., average [4] — 167 123 — — — (9)
Fem. [4] 350-380 158-187 100-120 — 1.4-1.8 (15) EGG-LAYING
Fem. [1] — 190 128 — — — (10)
Fem. [1] 375 164 126 — — — (9)
Unsexed [?] 375 200 138 115 — — (")
Unsexed [?] 355 220 130 110 — — (12) Place Months
Unsexed [?] 375 212 137 137 —
_ —
_ (13)
Unsexed [?] 375 200 138 115 (14) S Australia Aug-Jan
Unsexed [1] 360 195 128 — — — (8) N Australia Feb-May
Unsexed [1] 370 187 126 — — — (8) New Zealand (South Island) Nov-Dec
Unsexed [1] — — — 1.70 (5) W Java (formerly) Mar-Sep
—
Unsexed [1] — — — — - 1.68 (5)
Unsexed [1] — — - 1.29 (5)
— — _
Unsexed [1] —
—
— - 1.726 (6)
Unsexed [1] — — — — - 1.80 (7)
a
From specimen label.
References: (1) Ogilvie-Grant 1889. (2) Pers. obs. M.P.K. (British Mus./lO Apr 89). (3) Vestjens 1976:56. (4)
Pers. obs. M.P.K. (S. Austr. Mus./lO Apr 85). (5) B.P. Hall 1974:46. (6) Lavery 1971:52. (7) Serventy and
Whittell 1962:125. (8) Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (9) Pers. obs. M.P.K. (CSIRO, Canberra/4
Mar 85). (10) Pers. obs. M.P.K. (W. Austr. Mus./24 Apr 85). (11) Lucas and LeSouef 1911:126. (12) Rand and
Gilliard 1967:49. (13) Gould 1865:288. (14) Sharpe 1898:48. (15) K.W. Lowe (pers. comm., 4 Mar 90).

324
 Blackfaced Spoonbill
(Platalea minor)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [2] 338-360 173-180 118-125 _ _ _ (1)

Male, average [2] 249 176 121 _ _ _ (1)
Male [1] 365 180 119 105 - - (2)
Male [1] 362 180 125 _ _ _ (10)
Fern. [1] 376 188 135 _ _ _ (10)
Fern. [1] 350 180 125 _ _ _ (1)
Fern. [1] 350 183
_ 118 100 (2)
Fern, (imm.) [1] — — - - 1.338 (13)
Both [?] 356-371 183-185 119-121 102-107 (3)
Both [?] 338-376 — — 105-116 - - W
Both [?] 350-370 190-193 — _ _ _ (5)
Both [?] 334-358 151-196 106-115 103-122 (6)
Both [?] — — 108-134 _ _ _ (7)
Both [?] 338-371 145-193 119-134 100-117 - - (8)
Both [4] 338-369 145-163 108-133 105-113 - - (9)
Unsexed [1] 350 193 125 _ _ _ (1)
Unsexed [1] — 155 — _ _ _ (1)
Unsexed [1] 367 192 126 _ _ _ (10)
Unsexed [1] 366 178 125 — — — (10)
Unsexed [1] 361 190 132 _ _ _ (10)
Unsexed [1] 367 191 131 _ _ _ (11)
Unsexed [1] 361 178 120 101 - - (12)

References: (1) Ogilvie-Grant 1889. (2) Sharpe 1898:51. (3) Hackisuka 1931-32:347. (4) Hackisuka and
Udagawa 1951:102. (5) Hartert 1912-21:1220. (6) H.G. Won 1966. (7) B.F. King and E.G. Dickinson 1975:54.
(8) Kobayashi 1956b:67. (9) La Touche 1931-34:433. (10) Pers. obs. M.P.K. (British Mus./lO Apr 89). (11)
Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (12) duPont 1971:23. (13) Fennell and King 1964:241.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
Korea Jun-Jul
Extremes 10 58.2-66.0 40.4-48.0 (1)
Average 10 62.5 44.6 — (1)
Calculated 10 — — 68 (1)
Extremes — 63-67 41-42 — (2)
Average 10 63.7 42.6 — (3)
References: (1) Schonwetter 1967. (2) Won 1966. (3) Etchecopar and Hue
1978:80.

325
 African Spoonbill
(Platalea alba)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n] (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [?] 378-405 (1)

Male [1] 405 226 166 137 — (2)
Fern. [?] 363-380 — — — — — (1)
Fern. [?] 363-380 160-184 130-139 130-136 _ — (2)
Fern. [1] 370 175 135 128 - — (5)
Fern. [1] - — — — — 1.79 (4)
Both [?] 170-226 130-160 135-137 - — (1)
Both [?] 370-415 — — — — — (3)
Both [15] 365-414 172-230 131-157 105-152 - — (4)
Both, average [15] 384 193 144 124 - — (4)
Unsexed [1] - — — — — 1.59 (6)

Longevity in wild = 11 years 5 months (McLachlan and Liversidge 1978:57).

References: (1) Bouet 1955a:132. (2) Bannerman 1930:124. (3) Clancey 1964a:51. (4) Brown et al. 1982:208.
(5) Sharpe 1898:50. (6) FJ.Jackson 1938:88.

EGG M E A S U R E M E N T S EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
Senegal Oct-Mar
Extremes 23 59.1-72.2 40.6-48.0 (1) Guinea-Bissau Jan-Feb, May, Nov-Dec
Average 23 65.4 44.0 — (1) Sierra Leone Sep
Calculated 23 — — 69 (1) Niger Oct
Extremes 205 57-82 40-48 — (2) Mali Jan-Apr, Aug-Oct
Average 205 68 45 69 (2) Chad Jan-Apr
Extremes 15 65.2-73.9 40.5-47.2 — (3) Nigeria Jan-Feb
Average 15 69.1 44.5 — (3) Sudan Feb-Mar, Sep
Extremes 33 61.0-71.4 39.7-47.8 — (4) Ethiopia Jul-Aug, Oct
Average 33 66.4 44.2 — (4) Kenya Dec-Jul, Sep-Oct (peak in May)
Extremes 72 54.6-72.4 35.5-47.5 — (5) Uganda Feb-Jun
Average 72 66.2 44.5 — (5) Burundi Jun-Oct
Extremes 4 67-73 44-46 — (6) Tanzania Feb— Mar, May
Average 10 66.8 43.4 — (6) Zambia Feb-Jul
Extremes — 65-70 42-44 — (7) Malawi Sep
Zimbabwe Jan-Sep
References: (1) Schonwetter 1967. (2) Brown et al. 1982:209. (3) Botswana Mar-Jun
Broekhuysen 1957. (4) McLachlan and Liversidge 1970:57. (5) Whitelaw 1968. Namibia Feb-Aug, Oct
(6) FJ. Jackson 1938:90. (7) Bouet 1955a:132. South Africa (Cape Prov.) Aug—Nov
South Africa (Transvaal, O.F.S.) Mar-Sep
South Africa (Natal) Feb-Nov
Madagascar Feb

326
 Yellowbilled Spoonbill
(Platalea flavipes)

BODY MEASUREMENTS

Wing Culmen Tarsus Tail Wing-Span Weight

Sex W (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [5] 225-238 (i)

Male, average
Male
[5] — 231 — —
—
— •
—
—
1.75-2.0a
(i)
[3] 410-420 217-236 133-140 (2)
Male, average [3] 415 227 136 — — 1.92a (2)
Male, average [?] 400 213 133 175 — — (3)
Male, average [?] 413 225 138 170 — — (4)
Male [1] — — — 175 — — (2)
Male [1] —
_ 222 130 — 1375a 1.82a (6)
Male [1] 222 134 — — — (6)
Fern. [8] 186-231 — — — — — (1)
Fern., average [8] 205 — — — — — (1)
Fern. [4] — 179-191 113-122 — — — (2)
Fern., average M — 185 118 — — — (2)
Fern., average [?] 370 — — — — — (4)
Fern. [1] 395 — — 172 — 1.5a (2)
Fern. [1] — 195 119 — 1340a 1.7a (6)
Fern. [1] — 205 115 — — — (6)
Both, average [?] 413 225 138 170 — — (5)
Unsexed [1] — — — — 1450 1.8 (7)
Unsexed [1] 363 181 119 138 — — (8)
a
From specimen labels.
References: (1) Vestjens 1975a. (2) Pers. obs. M.P.K. (CSIRO, Canberra/4 Mar 85). (3) AJ. North 1913-
14:14. (4) Sharpe 1898:52. (5) Lucas and Le Souef 1911:127. (6) Pers. obs. M.P.K. (S. Austr. Mus./lO Apr 85).
(7) Serventy and Whittell 1962:126. (8) Gould 1865:289.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
S Australia Aug-Jan
Extremes 27 65.0-78.2 44.0-49.0 (i) N Australia Mar-Jun
Average 27 69.6 46.0 — (i)
Calculated
Extremes
27
13
—
65.5-77.0
—
43.3-47.5
81
—
(i)
(2)
Average 13 70.1 45.2 — (2)
Average 12 69 47 — (3)
Average 8 68 45 — (4)
Average — 70 47 — (5)
Average — 68 45 — (6)
Average — 60[?] 47 — (7)
References: (1) Schonwetter 1967 (2) AJ. North 1913-14:18. (3) Vestjens
1977b:56. (4) Serventy and Whittell 1962:126. (5) Lucas and Le Souef
1911:127. (6) Beruldsen 1980:159. (7) Frith 1976:91.
327
 Roseate Spoonbill
(Platalea ajaja)

BODY M E A S U R E M E N T S

Wing Culmen Tarsus Tail Wing-Span Weight

Sex [n]- (mm) (mm) (mm) (mm) (mm) (kg) Reference

Male [13] 345-375 156-173 107-112 93-107 (1)

Male, average [13] 356 166 109 99 — — (1)
Male [?] - — — — 'to about 'up to 1.6' (2)
1300'
Male [?] - — — — — 1.2-1.8 (3)
Male [?] 341-369 154-179 107-124 94-103 — — (8)
Male, average [?] 351 168 114 98 — — (8)
Male [1] 350 — — — — — (4)
Male [1] 345 163 105 93 — — (5)
Fern. [11] 325-365 145-177 97-117 87-108 — — (1)
Fern., average [11] 343 157 104 97 — — (1)
Fern. [?] — — — — 1.4-1.7 (3)
Fern. [?] 322-352 151-162 97-107 85-100 — — (8)
Fern, average •[?] 335 156 104 94 — — (8)
Fern. [1] 345 — — — — — (4)
Fern. [1] 333 — — — — — (5)
Both [?] 150-180 107-123 — — — (6)
Both [?] 360^370 163-172 — 97-105 — — (7)
Both, average [?] — — — 1275 1.6 (8)

Longevity in captivity = 10 years 5 months (Terres 1980:554).

References: (1) Blake 1977:202. (2) Palmer 1962:534. (3) Haverschmidt 1968:30. (4) ffrench 1973:78. (5)
Sharpe 1898:54. (6) J.C. Lewis 1983:1. (7) Goodalletal. 1951:108. (8) Oberholser and Kincaid 1974:132-133.

EGG MEASUREMENTS EGG-LAYING

Length Width Weight Place Months

No. (mm) (mm) (g) Reference
E Texas, SW Louisiana Apr-Jun
Extremes 50 59.5-71.0 41.2-46.5 (1) Florida Bay Nov-Dec
Average 50 65.2 43.8 — (1) Peninsular Florida Mar-Apr
Calculated 50 — — 68 (1) Mexico (Gulf coast) Feb
Extremes 70 56.9-71.2 41.3-47.9 52.4-77.0 (2) Honduras Feb-Mar
Average 70 64.4 43.7 62.1 (2) Cuba (Camaguey Prov.) Sep-Nov
Extremes 40 60.2-71.5 41.0-47.0 — (3) Surinam Aug-Sep
Average 40 65.0 43.9 — (3) Argentina (Buenos Aires Prov.) Oct-Dec
Average 20 65.0 44.2 — (4)
Average — 64.3 43.3 — (5)
Average — 65.0 43.7 — (6)

References: (1) Schonwetter 1967. (2) D.H. White et al. 1982. (3) Bent
1926:16. (4) Palmer 1962:538. (5) Sprunt 1954:49. (6) Oberholser and Kincaid
1974:133.

328

The following bibliography is based extensively on a list of references kindly supplied by Malcolm Coulter and the ICBP Storks, Ibises and
Spoonbills Group.
Abbott, C.G. 1930. Wood Ibises summering in
San Diego County, California. Condor 33:29-
30.
Abbott, C.G. 1934. Another invasion of Wood
Ibis in southern California. Condor 37:35-36.
Abbott, C.G. 1938. An exceptional influx of
Wood Ibises into south California. Condor
40:257.
Abdulali, H. 1967. On the occurrence of the
Black-necked Stork [Xenorhynchus asiaticus
(Latham)] in the Bombay Konkan./. Bombay
Nat. Hist. Soc. 64:367.
Abdulali, H. 1968. A catalogue of the birds in the
collection of the Bombay Natural History
Society, I (Gaviiformes toCiconiiformes)./.
Bombay Nat. Hist. Soc. 65:182-199.
D'Abreu, E.A. 1912. Notes on a bird collecting
trip in the Balaghat District of the Central
Provinces./ Bombay Nat. Hist. Soc. 21:1158-
1169.
Adamiak, W. 1974. Bocian czarny Ciconia nigra w
powiecie Jarocin wojewodztwa poznanskiego.
Chronmy Przyr. Ojczysta 29:68-70.
Adamian, M.S. 1991. European White Storks
and other Ciconiiformes in the Armenian
Republic. Spec. Group Storks, Ibises, Spoonbills
Newsletter 4(l):2-3.
Adams, A.L. 1858. Notes on the habits, haunts,
etc. of some of the birds of India. Proc. Zoo/.
Soc. Lond. 1858:466-512.
Adams, D.A. 1963. Battery Island 1963. Chat
27:65-68.
Agoramoorthy, G., and S.M. Mohnot. 1989.
Checklist of birds around Jodhpur. Tigerpaper
16:11-13.
Aguilera, E. 1989. Sperm competition and
copulation intervals of the White Spoonbill
(Platalea leucorodia, Aves, Threskiornithidae).
Ethology 82:230-237.
Aguilera, E. 1990a. Parental infanticide by White
Spoonbills Platalea leucorodia. Ibis 132:124-125.
Aguilera, E. 1990b. Sexual differences in nest
attendance and chick-feeding rhythms of
White Spoonbills. Auk 107:416-420.
Aguilera, E. in press. Evolutionary consequences
of sperm competition on parental roles in the
White Spoonbill (Platalea leucorodia). Behaviour.
Aguilera, E., and F. Alvarez. 1989. Copulations
and mate guarding of the Spoonbill (Platalea
leucorodia). Behaviour 110:1-22.
Aguilera, E., and J. Sanudo. 1986. Pasado y
presente de las colonias de Ciconiformes en
Donana y Odiel. Bios 2:11-13.
Aharoni, J. 1911. An den Brutplatzen von
Comatibis comata. Ehrbg. Z. Ool. (Stuttg.) 1(2):9-
11.
Aharoni, J. 1912. Houbara macqueeni Gray. Orn.
Jahb. 23:1-15.
Aharoni, J. 1929. Zur Brutbiologie von Comatibis
comata Bp. (Eudocimus eremita L.). Beitr.
Fortpflanzungsbiol. Vogel 5:17-19.
Aharoni, J. 1932. Bemerkungen und
Erganzungen zu R. Meinertzhages Werk
'Nicoll's Birds of Egypt'. J. Orn. 80:416-424.
Akcakaya, H.R. 1990. Bald Ibis Geronticus eremita
population in Turkey: an evaluation of the
captive breeding project for reintroduction.
Biol. Conserv. 51:225-237.
Akester, A.R., D.E. Pomeroy, and M.D. Purton.
Bibliography
1973. Subcutaneous air pouches in the
Marabou Stork (Leptoptilos crumeniferus). J.
Zoo/. (Lond.) 170:493^99.
Akhtar, S.A. 1947. Ab-Istadeh, a breeding place
of the flamingo (Phoenicopterus ruber roseus
[Pallas]) in Afghanistan./. Bombay Nat. Hist.
Soc. 47:308-314.
Alamargot, J. 1976. La nidification du Marabout
d'Afrique (Leptoptilos crumeniferus) en colonie
dans le sud-ouest de L'Ethiopie. Oiseau Rev. Fr.
Orn. 46:178-182.
Alamargot, J. 1984. New nesting colony of
Marabou Storks (Leptoptilos crumeniferus).
Ethiopian Wildl. Nat. Hist. Soc. Newsletter 188:2.
Ali, M.M. 1971. A new species of dioctophimid
nematode, Eustrongylides indicus n. sp., from an
Indian bird. Riv. Parasitol. 32:47-50.
Ali, S. 1934. The Hyderabad State ornithological
survey, part 5. J. Bombay Nat. Hist. Soc. 37:425-
454.
Ali, S. 1945. The Birds ofKutch. Oxford
University Press, London. 175pp.
AH, S. 1953. The Birds ofTravancore and Cochin.
Oxford University Press, London.
Ali, S. 1954. The birds of Gujarat, part \.J.
Bombay Nat. Hist. Soc. 52:384-458.
Ali, S. 1964. The Book of Indian Birds, 7th edn.
Bombay Natural History Society, Bombay.
158pp.
Ali, S. 1979. The Book of Indian Birds. Bombay
Natural History Society, Bombay.
Ali, S. 1981. Studies on the movement and
population structure of Indian avifauna.
Bombay Nat. Hist. Soc. Ann. Rep., 1 (1980-1981).
Ali, S., and S.D. Ripley. 1968. Handbook of the
Birds of India and Pakistan, vol. 1. Oxford
University Press, London.
Ali, S., and S.D. Ripley. 1983. A Pictorial Guide to
the Birds of the Indian Subcontinent. Bombay
Natural History Society, Oxford University
Press, Oxford.
Allan, D. 1982. The status of the Bald Ibis in the
Transvaal, 1982 progress report. Unpublished
report for Nature Conservation Division,
Transvaal Provincial Administration, South
Africa.
Allan, D. 1983a. The status of the Bald Ibis in
the Transvaal, 1983 progress report.
Unpublished report for Nature Conservation
Division, Transvaal Provincial
Administration, South Africa.
Allan, D. 1983b. Breeding success of the Bald
Ibis, 1983 progress report. Project TN 6/4/4/5.
Unpublished report for Nature Conservation
Division, Transvaal Provincial
Administration, South Africa.
Allan, D. 1984a. A study of coloniality,
adaptations to cliff-nesting, productivity,
breeding habits, vocalizations and parent-
young recognition in the Bald Ibis (Geronticus
calvus) in southern Africa. Project TN 6/4/4/5.
Unpublished report for Nature Conservation
Division, Transvaal Provincial
Administration, South Africa.
Allan, D. 1984b. Southern African White Stork
census. Bokmakierie 36:77-80.
Allan, D. 1985a. White Stork census results
1984/85. Afr. Wildl. 39:251-252.
Allan, D. 1985b. The national stork counting
329
weekend 18-20 January 1985. Bokmakierie
37:112-113.
Allan, D. 1985c. The Bald Ibis-a false alarm?
Fauna Flora (Transv.) 42:16-21.
Allan, D. 1989a. The Bald Ibis returns to Cape
Town. Quagga 25:18-20.
Allan, D. 1989b. Results of a national stork
counting weekend in South Africa. Pp. 365-
372 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Allavena, S. 1976. Una Cicogna nera (Ciconia
nigra) al Lago Barrea nel Parco Nazionale
d'Abruzzo. Riv. Ital. Orn. 46:102-110.
Allen, J.A. 1876. Occurrence of the Wood Ibis in
Pennsylvania and New York. Nuttall Orn. Club
Bull. 1:96.
Allen, J.A. 1908. The generic names Mycteria and
Tantalus of Linnaeaus, 1758. Auk 25:37-38.
Allen, R.P. 1935. Notes on the Roseate Spoonbill
on the Gulf Coast. Auk 52:77-78.
Allen, R.P. 1936. With the Audubon wardens
recent news of the great white heron, glossy
ibis and other rare species. Bird Lore 38:295-
297.
Allen, R.P. 1938. The sanctuary front: condition
of the Great White Heron and Spoonbills;
progress and obstacles in other fields. Bird Lore
40:125-129.
Allen, R.P. 1942. The Roseate Spoonbill.
Research report 2, National Audubon Society,
New York, New York.
Allen, R.P. 1947. The Flame Birds. Dodd, Mead
and Co., New York. 233pp.
Allen, R.P. 1955. The White Ibis situation in
Louisiana. Louisiana Conserv. 7(5): 17-19.
Allen, R.P. 1957. On the Trail of Vanishing Birds.
McGraw-Hill, New York.
Allen, R.P. 1958. A progress report on the
wading bird survey. Unpublished report,
National Audubon Society.
Allen, R.P. 1962. Roseate spoonbills radiant
birds of the Gulf Coast. Natl Geographic
121:274-288.
Allen, R.P. 1963. The White Ibis Situation in
Louisiana. Louisiana Wildlife and Fisheries
Commission, New Orleans.
Allen, R.P. 1964. Our only native stork, the
Wood Ibis. Natl Geographic 125:294-306.
Allen-Grimes, A.W. 1982. Breeding biology of
the White Ibis (Eudocimus albus) at Battery
Island, North Carolina. M.S. Thesis,
University of North Carolina, Wilmington,
North Carolina.
Allouse, B.E. 1953. The Avifauna of Iraq. Iraq
Nat. Hist. Mus. Publ. 3:1-164.
Allport, G., and S. Wilson. 1986. Results of a
census of the Milky Stork in West Java. Report of
the Indonesian Waterbird Survey Expedition,
July-October 1984. ICBP report 14,
Cambridge, England.
Al-Safadi, M.M. 1990. Observations on the
breeding of the Abdim's Stork Ciconia abdimii
in Yemen. Sandgrouse 12:43-52.
Amadon, D. 1953. Avian systematics and
evolution in the Gulf of Guinea. Bull. Am. Mus.
Nat. Hist. 100:395-451.
Amadon, D. 1966. The superspecies concept.
System. ZooL 15:245-249.
Amadon, D., and G. Woolfenden. 1952. Notes on
the Mathew's collection of Australian birds:
The order Ciconiiformes, Am. Mus. Novit.
1564:1-16.
Ambedkar, V.C. 1959. The occurrence of the
White-necked Stork [Ciconia episcopus
(Boddaert)] in the Kashmir Valley. J. Bombay
Nat. Hist. Soc. 56:633,
American Ornithologists' Union. 1908. 14th
supplement to the A.O.U. Checklist. Auk
25:343-399.
American Ornithologists' Union. 1957. The AOU
Check-list, 5th edn. Lord Baltimore Press,
Baltimore, Maryland.
American Ornithologists' Union. 1983. Check-list
of North American Birds, 6th edn. Allen Press,
Lawrence, Kansas.
Amget, B. 1986. Some Information on the Openbilled
Stork. Wildlife Conservation Division, Royal
Forest Department, Bangkok, Thailand.
Anderson, A.B. 1949. Marabou nesting colonies
of the southern Sudan. Sudan Notes Rec. 30:114—
118.
Anderson, D.J., and RJ. Horowitz. 1979.
Competitive interactions among vultures and
their avian competitors. Ibis 121:505-509.
Anderson, J.M. 1978. Protection and
management of wading birds. Pp. 99—103 in
Sprunt, A., IV, J.C. Ogden, and S. Winckler
(eds), Wading Birds. Research report 7,
National Audubon Society, New York.
Andersson, C J. 1872. The Birds ofDamara Land.
London.
Anmuller, S. 1952. Der Bestand an Weiss- und
Schwarzstorchen in Burgenland im Jahre
\95l.HeimatbL 14:79-91.
Anonymous. 1931. Indo-Chinese Birds. Field
Mus. Nat. Hist. Publ. ZooL Ser. 290:18.
Anonymous. 1951. A breeding record of the
Glossy Ibis. Ostrich 22:108-110.
Anonymous. 1955. Florida birdlife—Eastern
Glossy Ibis. Florida WildL 8(12):5.
Anonymous. 1957. Florida birdlife—ibis,
spoonbill, heron, and cattle egret. Florida
WildL ll(2):41-42.
Anonymous. 1968. The Japanese Crested Ibis or
Toki. Biol. Conserv. 1:36.
Anonymous. 1969. Storks, ducks, raptors, crakes,
and waders. British Birds 62:87-88.
Anonymous. 1970. Four new entries in the little
red book. New Sci. 47:457.
Anonymous. 1973. The Bald Ibis at Bireck. Bull.
Orn. Soc. Turkey 9:6-7.
Anonymous. 1974a. Remains of flightless ibis
found in Hawaiian Islands. Inform. Bull. Pacific
Sci.Assoc. 26:10.
Anonymous. 1974b. Glossy Ibis nesting in
southern Florida. Fla. Field Nat. 2:13-14.
Anonymous. 1980. Natal Bald Ibis survey. Natal
Wildl.,]u\y 1980.
Anonymous. 1983. Jabiru nesting in Belize. Belize
Audubon Soc. Bull. 15:1-2.
Anonymous. 1984. Scarlet Ibis: model species for
conservation. Flying Free 2:3-4.
Anonymous. 1985a. One of the finest sights at
the zoo. Zoorama Malaysia 85(1,2):!.
Anonymous. 1985b. Regional roundup. Interwader
Newsletter 5:1-7'.
Anonymous. 1985c. White Stork hangs on.
Newsreview, February 9.
Anonymous. 1986. Regional reports—Indonesia.
Oriental Bird Club Bull. 3:35.
Anonymous. 1987a. Recent reports: China.
Oriental Bird Club Bull. 5:33.
Anonymous. 1987b. Recent reports: Malaysia.
Oriental Bird Club Bull. 5:34.
Anonymous. 1988a. Nest sites of Black Storks
(Ciconia nigra) in Lasy Janowskie, Prov.,
Tarnobrzeg. Notatki Orn. 1988:227-231.
Anonymous. 1988b. Wood Storks in Georgia.
Nongame Wildlife Program pamphlet, Georgia
Department of Natural Resources, Atlanta,
Georgia. 8pp.
Anonymous. 1989. Milky Stork colonies located
in Sumatra. World Birdwatch 11(1):3.
Anonymous. 1990a. Estado de las poblaciones
del Corocoro rojo (Eudocimus ruber) en
Venezuela: diagnostico, conclusiones, y
recommendaciones para un plan de
conservacion. Pp. 137-151 in Frederick, P.C.,
L.G. Morales, A.L. Spaans, and C.S. Luthin
(eds), The Scarlet Ibis (Eudocimus ruber):
Status, Conservation, and Recent Research, Proc. 1st
International Scarlet Ibis Conserv. Workshop,
Caracas, Venezuela. IWRB Special
Publication 11, Slimbridge, Glos, UK.
Anonymous. 1990b. News from Wat Pai Lorn—
the Thailand Openbill Stork colony. Flying
FreeS:7-9.
Anonymous. 199la. The SIS Group, Indian
survey of storks, and SIS Newsletter. Zoos'
Print 6(l)'A.
Anonymous. 1991b. New Zealand's population
of Royal Spoonbills on the increase (from
RAOUNew Newsletter no. 84, June 1990). Spec.
Group Storks, Ibises, Spoonbills Newsletter 4(1):7.
Anonymous. 1991c. Adjutant Stork survives in
Assam. Oryx 25:69.
Anonymous. 199Id. More exciting news from
Vietnam. World Birdwatch 13(3):4.
Antas, P.T.Z. 1979. Breeding the Scarlet Ibis
Eudocimus ruber at the Rio de Janeiro Zoo. Int.
ZooYb. 19:135-139.
Antas, P.T.Z., and R.B. Cavalcanti. 1988. Aves
Comuns do Planalto Central. Editora
Universidade de Brasilia, Brasilia.
Antas, P.T.Z., P. Roth, and R.I.G. Morrison.
1990. Status and conservation of the Scarlet
Ibis (Eudocimus ruber) in Brasil. Pp. 130-136 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conserv. Workshop, Caracas, Venezuela.
IWRB Special Publication 11, Slimbridge,
Glos, UK.
Anthony, AJ. 1977. A further breeding record of
the Woolly-necked Stork. Honeyguide 89:36-40.
Anthony, AJ. 1978a. Hadadah nesting at
Chipinda Pools, Gonarezhou National Park.
Honeyguide 95:33-35.
Anthony, AJ. 1978b. A further breeding record
of the Woolly-necked Stork: the 1977/78
season. Honeyguide 93:45.
Anthony, AJ., and B.Y. Sherry. 1980. Openbill
Storks breeding in the southeastern lowveld of
Zimbabwe Rhodesia. Ostrich 51:1-6.
Antipa, C., E. Girjabu, R. Iftimovici, and N.
Draganescu. 1984. Seriological investigations
concerning the presence of antibodies to
arboviruses in wild birds. Rev. Roum. Med.
Virol. 35:5-10.
Appert, O. 1966. Beitrag zur Biologic und zur
Kenntnis der Verbreitung des, Madagaskar
Mahnenibisses, Lophotibis cristata (Boddaert).
J. Orn. 107:315-322.
Arballo, E. 1990. Nest of Theristicus caudatus in
Uruguay. Romero 13:164-166.
Archer, A.L. 1978. Marabous breeding at
Kiboko, Kenya. East Afr. Nat. Hist. Soc. Bull.
(Sept./Oct.):106-107.
Archer, G., and E.M. Goodman. 1937. The Birds
of British Somaliland and the Gulf of Aden, vol. 1.
Gurney and Jackson, London. 285pp.
Archibald, G.W. 1978. Crane and ibis
conservation Republic of Korea. In Jackson, P.
(ed.), WWFYb: 1977-78.
Archibald, G. 1981. They've captured the Toki!
Int. WildL 11:20-24.
Archibald, G., and S. Lands. 1979. Conservation
of the Japanese Crested Ibis. Proc. Colonial
Waterbird Group (1978):1-15.
330
Archibald, G.W., S.D.H. Lands, L.R. Lands,
and I. Munetchika. 1980. Endangered ibises
(Threskiornithinae): their future in the wild
and in captivity. Int. Zoo Yb. 20:6-17.
Archibald, K.M. 1985a. Studbook of the Eastern
White Stork (Ciconia bqyciana) in captivity.
Baraboo, Wisconsin, USA.
Archibald, K.M. 1985b. Captive management of
the Eastern White Stork, Ciconia boyciana. Avic.
Mag. 91:92-101.
Archibald, K. 1991. Recommendations for
captive management of the Oriental White
Stork (Ciconia bqyciana). Pp. 193-202 in
Coulter, M.C., Q. Wang, and C.S. Luthin
(eds), Biology and Conservation of the Oriental
White Stork (Ciconia Boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Archibald, K., and B. Schmitt. 1991.
Comparisons between the Oriental White
Stork Ciconia c. boyciana and the European
White Stork Ciconia c. ciconia. Pp. 3-37 in
Coulter, M.C., Q. Wang, and C.S. Luthin
(eds), Biology and Conservation of the Oriental
White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Armstrong, E.A. 1965. Bird Display and Behaviour,
revised edn. Dover Publications, New York.
Armstrong, J.T. 1965. Breeding home range in
the nighthawk and other birds: its evolutionary
and ecological significance. Ecology 46:619-
629.
Arndt, R. 1974. The cliff dwellers. InL WildL
4:18-20.
Arnold, K.A. 1978. A Jabiru (Jabiru mycteria)
specimen from Texas. Auk 95:611-612.
Arnold, P. 1990. Reintroduction de la Cigogne
Blanche (Ciconia ciconia) en Alsace. Rev. Ecol.
Terre Vie Suppl. 5:253-256.
Arthur, S.C. 1931. The Birds of Louisiana. Bull. 20,
Louisiana Department of Conservation, Baton
Rouge, Louisiana, USA.
Aschenbrenner, L., and H. Schifter. 1975. Der
Bestand des Weiss-Storches (Ciconia ciconia L.)
in Oesterreich im Jahre 1974. Egretta 18(1):8-
17.
Ash, J.S. 1981. Bird-ringing results and ringed
bird recoveries in Ethiopia. Scopus 5:85-101.
Ash, J.S. 1989. The status of the White Stork in
Ethiopia, Somalia and Djibouti. Pp. 341-353
in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weiss-Storch: Status undSchutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Ash, J.S., and T.R. Howell. 1977. The Bald Ibis
or Waldrapp Geronticus eremita in Ethiopia.
Bull. Br. Orn. Club 97:104.
Ash, J.S., and J.E. Miskell. 1983. Birds of
Somalia, their habitat, status and distribution.
Scopus spec, suppl. 1. 97 pp.
Ash, J.S., RJ. Dowsett, and F. Dowsett-Lemaire.
In press. Additions to the East African
avifauna. Scopus.
Asian Wetland Bureau. 1988. Milky stork
colonies discovered. Asian Wetland News
1(2):12.
Asian Wetland Bureau. 1989a. Captive breeding
programme for Milky Storks. Asian Wetland
News 2(1):5.
Asian Wetland Bureau. 1989b. Storm's Stork
discovered breeding in Sumatra. Asian Wetland
Afow2(l):14.
Asian Wetland Bureau. 1990a. Malaysia's Milky
Storks breed again. Asian Wetland News 2(2): 1.
Asian Wetland Bureau. 1990b. North-west
Sichuan peat mining likely to threaten rare
waterbirds. Asian Wetland News 2(2):6.
Assfalg, W. 1979. Wenn dem Storch das
Klappern vergeht. Magazin Schwab. Zeitung zeil
weit 15 June.
Assfalg, W. 1980. Storchenburg von Mengen.
Schwab. Heimat 34:278-286 (Erganzung S.275).
Assfalg, W. 1982. Storche in Oberschwaben, die
letzten in Wurtternberg. Saulgauer Hefte 2:35-
47.
Assfalg, W. 1983. Storchenkampfe in
Oberschwaben. Schwab. Heimat 34:120-124.
Assfalg, W. 1986. Storche in Oberschwaben,
Pendler zwischen Ost und West. Naturschutz
Bad. Wurtt. 43:305-314.
Assfalg, W. 1987. Die Storche. Pp. 209-210 in
Schmid, J. (ed.) Chronik der Germeinde
Oggelshausen, 2. Aufl. Federsee-Verlag.
Assfalg, W., and E. Schuz. 1988. WeiBstorch-
Dreiergruppe, ein moglicher Umweg zur
Paarbildung. Orn. Mitt. 40:279-286.
Audubon,JJ. 1835. Ornithological Biography<,
Volume 3. Black, Edinburgh.
Audubon,JJ. 1840. The Birds of America.
Audubon, New York.
Audubon,JJ. 1967. The Birds of America. Dover
Publications, New York, New York.
Aust. C.S.I.R.O. Div. Wildl. Res. 1969. An index
of Australian bird names. Aust. CSIRO Div.
Wildl. Res. Tech. Paper. 20:1-93.
Austin, O.L., Jr. 1948. The Birds of Korea. Bull.
Mus. Comp.Zool. 101:1-301.
Austin, O.L. 1961. Birds of the World. Golden
Press, New York. 361 pp.
Austin, O.L., Jr. 1967. The Birds of the Gainsville
Region, Then and Now. University of Florida
Press, Gainsville, Florida.
Austin, O.L., Jr., and N. Kuroda. 1953. The
birds of Japan, their status and distribution.
Bull. Mus. Comp. Zoo/. 109:280-637.
Ayarzaguena Sanz, J., J.T. Perez, and C.H.
Ramo. 1981. Los Garceros del Llanos.
Cuadeernos Lagoven, Caracas, Venezuela.
Aymar, G.C. 1935. Bird Flight. Garden City,
New York.
Baal, F.L J. 1990. Coastal wetlands conservation
in Suriname. Pp. 71-76 in Frederick, P.C.,
L.G. Morales, A.L. Spaans, and C.S. Luthin
(eds), The Scarlet Ibis (Eudocimus ruber):
Status, Conservation, and Recent Research, Proc. 1 st
International Scarlet Ibis Conserv. Workshop,
Caracas, Venezuela. IWRB Special
Publication 11, Slimbridge, Glos, UK.
Babko, V.M. 1987. Nesting of the White Stork
(Ciconia ciconia) in the southwestern part of
Chernigov Region. Ornitologiya 22:174.
Babko, V.M. 1987. [On the records and breeding
of the Black Stork (Ciconia nigra) in the
southwestern part of Chernigov Region].
Ornitologiya ^\\1^.
Backhurst, G.C., P.L. Britton, and C.F. Mann.
1973. The less common Palaearctic migrant
birds of Kenya and Tanzania. J. E. Afr. Nat.
Hist. Soc. Nat. Mus. 140:1-38.
Badenhorst, M. 1956. The effect of a hailstorm
on wintering European storks. Ostrich 27:89-
90.
Badshah, M.A. 1963. Breeding colony of Painted
Storks. /Woe* 3:15-17.
Baerg, W.J. 1931. Birds of Arkansas. Bull. Ark.
Agric. Exp. Sta. No. 258.
Bailey, A.M., and R.J. Neidrach. 1937. Notes on
Colorado birds. Auk 54:524-527.
Bailey, R.F. 1934. New nesting records of Glossty
Ibis. Emu 33:279-291.
Bailey, W.L. 1875. Our Own Birds, a Familiar
Natural History of the Birds of the United States.
J.B. Lippincott, Philadelphia.
Bain, J.R., and S.R. Humphrey. 1980. Pp. 173-
196 in A Profile of the Endangered Species of
Thailand. Report 4, Florida State Museum
Office of Ecological Services, Tallahassee,
Florida, USA.
Baird, S.F., T.M. Brewer, and R. Ridgway.
1884. The Waterbirds of North America. Memoirs of
the Museum of Comparative Zoology 12. Little,
Brown and Company, Boston.
Bairlein, F. 1979. Zum Zugbeginnjunger Weiss-
Storche (Ciconia ciconia) in
Suedwestdeutschland. [On the onset of
migration of Juvenal White Storks (Ciconia
ciconia) in South-West Germany.] Vogelwarte
30(l):6&-69.
Bairlein, F. 1981. Analyse der Ringfunde von
Weiss-Storchen (Ciconia ciconia) aus
Mitteleuropa westlich der Zugscheide: Zug,
Winterquartier, Sommerverbreitung.
Vogelwarte 31:14-32.
Bairlein, F. in press. Population studies of White
Storks (Ciconia ciconia) in Europe, with special
reference to the western population. Proc.
Symp. Population Studies in Birds: Their Relevance
for Conservation and Management, Tour du Valet,
1988.
Bairlein, F., and G. Zink. 1979. Der Bestand des
Weiss-Storchs (Ciconia ciconia) in
Suedwestdeutschland: Eine Analyse der
Bestandsentwicklung. [The population of the
white stork (Ciconia ciconia) in South-West
Germany: an analysis of the population
dynamics.]/ Orn. 120(1): 1-11.
Baker, E.C.S. 1899. The birds of North Cachar.
Part X.J. Bombay Nat. Hist. Soc. 12:486-510.
Baker, E.C.S. 1929. The Fauna of British India,
including Ceylon and Burma. Birds, vol. VI.
Taylor and Francis, London.
Baker, E.C.S. 1930. The Fauna of British India,
including Ceylon and Burma. Birds, vol. VIII.
Taylor and Francis, London, England.
Baker, E.C.S. 1935. The Nidification of Birds of the
Indian Empire, vol. IV. Taylor and Francis,
London. 546 pp.
Baker, G. 1982. Review of the status; U.S.
breeding populations of the Wood Stork.
Federal Register 47:6675-6677.
Baker, J.H. 1939. Counting spoonbills and
mosquitoes. Bird Lore 41:244.
Baker, J.H. 1951. Pinks are showing up. Audubon
Mag. 53:127.
Baker, W. 1850. On the capture, habits and
change of plumage in the Black Stork (Ciconia
nigra). Zoologist 8:2700-2703.
Bakewell, D.N., and L. Young. 1989. Report on
Ornithological Observations from Hangzhou Bay and
Yencheng Nature Reserve, East China. Publication
58, Asian Wetlands Bureau, Kuala Lumpur.
Baldwin, P.J., and B.S. Meadows. 1988. Birds of
Madinal Yanbu Al-Sinaiyah and its Hinterland.
Royal Commission for Judail and Yanbu,
Riyadh, Kingdom of Saudi Arabia.
Ball, V. 1878. From the Ganges to Godaveri.
Stray Feathers 7:191-235.
Ballance, D.K., and S.L.B. Lee. 1961. Notes on
autumn migration at the Bosphorus and in the
Aegean. Ibis 103a: 195-204.
Balogh, G. Nesting of the Black Stork (Ciconia
nigra) in the environs of Leninvaros. Aquila
87:141.
Balthasar, J., and K.J. Schafer. 1968.
Braunsichler (Plegadis falcinellus) in Westfalen.
Orn. Mitt. 20:45.
Baltzer, M.C. 1990. A report on the wetland
avifauna of South Sulawesi. Kukila 5:27-55.
Bamford, O.S., and G.M.O. Maloiy. 1980.
Energy metabolism and heart rate during
treadmill exercise in the Marabou Stork. J.
Appl. Physiol. 49:491-496.
Bancroft, G.T., J.C. Ogden, and B.W. Patty.
1988. Wading bird colony formation and
turnover relative to rainfall in the Corkscrew
Swamp area of Florida during 1982-1985.
Wilson Bull. 100:50-59.
Banerjee, A.K. 1984. A brief account of breeding
activities of large waders with special emphasis
on Ciconiformes in Saurashtra Region,
Gujarat, India. Unpublished report,
Saurashtra University, India.
Banerjee, D.P., S.P. Baudekar and V.K.
Paralkan. 1990. Aggressive behaviour of
331
Blacknecked Storks toward Cranes. J. Bombay
Nat. Hist. Soc. 87:140.
Banet, L. 1975. Storks and ospreys passing by
Bazillac. Alauda 42:503.
Bangs, O. 1918. Vertebrata from Madagascar-3.
Aves. Bull. Mus. Comp. Zool. 61:489-511.
Bangs, O., and T.E. Penard. 1918. Notes on a
collection of Surinam birds. Bull. Mus. Comp.
Zool. 62:23-93.
Bangs, O., and J.L. Peters. 1928. Birds collected
by Dr. Joseph F. Rock in western Kansu and
eastern Tibet. Bull. Mus. Comp. Zool. 68(1):324.
Bangs, O., and J. Van Tyne. 1931. Birds of the
Kelley-Roosevelt expedition to French Indo-
China. Field Mus. Nat. Hist. Publ. Zool. Ser.
18:39-40.
Bangs, O., and W.R. Zappey. 1905. Birds of the
Isle of Pines. Amer. Nat. 39:179-215.
Banner, A. 1989. A habitat suitability model for
the Wood Stork in southern Florida. Colonial
Waterbird Soc. Newsletter 13(2): 17 (abstr.).
Bannerman, D.A. 1930. The Birds of Tropical West
Africa, vol. 1. Oliver and Boyd, London.
Bannerman, D.A. 1953. The Birds of West and
Equatorial Africa, vol. 1. Oliver and Boyd,
London.795pp.
Bannerman, D.A. 1957. The Birds of British Isles,
vol. 6. Oliver and Boyd, Edinburgh. 326pp.
Bannerman, D.A. 1958. Birds of Cyprus. Oliver
and Boyd, London.
Bannerman, D.A. 1963. Birds of the Atlantic
Islands, vol. 1. A History of the Birds of the Canary
Islands and the Salvages. Oliver and Boyd,
Edinburgh. 358pp.
Bannerman, D., and J. Bannerman. 1953. A
second journey to the Moroccan Sahara (in
1952) and over the Great Atlas. Ibis 95:128-
139.
Bannerman, D.A., and M.A. Bannerman. 1965.
Birds of the Atlantic Islands, vol. 2. A History of the
Birds of Madeira, the Desertas, and the Porto Santo
Islands. Oliver and Boyd, Edinburgh. 206pp.
Bannerman, D.A., and W.M. Bannerman. 1966.
Birds of the Atlantic Islands, vol. 3. A History of the
Birds of the Azores. Oliver and Boyd,
Edinburgh. 262pp.
Bannerman, D.A., and W.M. Bannerman. 1968.
Birds of the Atlantic Islands, vol. 4. A History of the
Birds of the Cape Verde Islands. Oliver and Boyd,
Edinburgh. 458pp.
Barbour, D.J. 1968. New distributional data.
Ostrich 39:271.
Barbour, T. 1923. Birds of Cuba. Memoirs of the
Nuttall Ornithological Club 6, Cambridge,
Massachusetts.
Barbour, T. 1943. Cuban Ornithology. Memoirs of
the Nuttall Ornithological Club 9, Cambridge,
Massachusetts.
Bardin, P. 1959. Baguage de cigognes blanches,
C. ciconia, de 1950 a 1955, dans la region de
Souk-el-Arba (Tunis). Mem. Soc. Sci. Nat.
Tunisie. 4:95-101.
Baris, S. 1989a. Final report of the Bald Ibis Project.
Unpublished report (10 Sept 1989), The
Society for the Protection of-Nature, DHKD,
Turkey.
Baris, S. 1989b. Turkey's habitats and
ornithological importance—Bald Ibis.
Sandgrouse 11:48.
Barlow, C.S. 1932. An ornithological excursion
to Zululand. Ostrich 3:75-81.
Barlow, C.S. 1933. A wonderful nesting colony of
Sacred Ibis, herons, and other waders at
Johannesberg. Ostrich 4:5-9.
Barnes, C.T. 1919. Roseate Spoonbill in Utah.
Auk 36:565-566.
Barlow, J.C. 1966. Status of the Wood Ibis, the
Fulvous Tree Duck and the Wheatear in
Ontario. Can. Field-Nat. 80:183-186.
Barnes, P.R. 1950. Nesting of the Black Stork,
Ciconia nigra, in Natal. Ostrich 21:39-40.
Barrett, C. 1936. Glossy Ibis nesting in Victoria.
Victorian Nat. 52:169-171.
Barrett, C. 1947. Australian Bird Life. Geoffrey
Cumberlege, Oxford University Press,
Melbourne, Australia.
Barrett, C. 1954. Wild Life in Australia and New
Guinea. Heinemann, London.
Barros, P., de, and D. Moura. 1989. Breeding,
feeding and mortality factors in the White
Stork populations nesting in Portugal in 1985.
Pp. 405-414 in Rheinwald, G., J. Ogden, and
H. Schulz (eds.), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DBA, Braunschweig,
Germany.
Barrows, W.B. 1912. Michigan Bird Life.
Michigan Agricultural College, Lansing.
Barthos, G. 1955. White Stork injured by
hailstorm. Aquila 59-62:456.
Barthos, J. 1908. Kampf der Weissen Storche um
das Nest. Aquila 15:310.
Bartlett, A.D. 1860. Note on the Balaeniceps rex.
Proc. Zool. Soc. Land. 1860:461.
Bartlett, A.D. 1861. On the affinities of
Balaeniceps. Proc. Zool. Soc. Lond. 1861:131-134.
Bartlett, A.D. 1866. Notes on the breeding of
several species of birds in the Society's garden
during the year 1865. Proc. Zool. Soc. Lond.
1866:76-79.
Barus, V. 1966. Nematodos de la familia
Tropisuridae Yamaguti, 1961, parasites de
aves de Cuba. Poeyana 20:1-22.
Basil-Edwards, S. 1926. A contribution to the
ornithology of Dehli, part 2.J. Bombay Nat.
Hist. Soc. 31:567-578.
Basilio, C.M.F. 1963. Aves de la Isla de Fernando
Poo. Coculsa, Madrid.
Bassett, T.H. 1963. Japanese cranes and storks.
Can. Audubon 25:117-120.
Bateman, D.L. 1970. Movement-behavior in
three species of colonial nesting wading birds:
a radiotelemetric study. Ph.D. Dissertation,
Auburn University, Alabama.
Bates, R.S.P. 1924. Birds-nesting with a camera
in India./ Bombay Nat. Hist. Soc. 30:306-313.
Batten, T.A. 1943. Sacred Ibis at East London
(Threskiornis aethiopica aethiopica). Ostrich
14:116-117.
Bauer, K. 1952. Ausbreitung des
Schwarzstorches in Oesterreich. Vogelwelt
73:125-129.
Bauer, K. 1965. Entwicklung und Bestand der
oesterreichischen Vogelfauna; vorlaufiger
Versuch einer quantitativen Beurteilung. Nat.
Land 1:16-19.
Bauer, K., H. Freundl, and R. Lugitsch. 1955.
Weitere Beitrage zur Kenntnis der Vogelwelt
des Meusiedlersee-Begietes. Wiss. Arb.
Bungenland 7.
Bauer, K.M., and U.N. Glutz von Blotzheim.
1966. Handbuech der Voegel Mitteleuropas, Band 1.
Gaviiformes—Phoenicopteriformes. Akademische
Verlagsgesellschaft, Frankfurt am Main.
483pp.
Baynard, O.E. 1912. Food of herons and ibises.
Wilson Bull. 24:167-169.
Baynard, O.E. 1913a. Breeding birds of Alachua
Co., Fla. Auk 30:240-247.
Baynard, O.E. 1913b. Home life of the Glossy
Ibis (Plegadis autumnolis Linn.). Wilson Bull.
25:103-117.
Baynard, O.E. 1913-1914. Two months in the
Everglades. Oologist 30:287-294; 31:32-40.
Baynard, O.E. 1914a. The White Ibis. Bluebird
(St. Louis) 7:16-22.
Baynard, O.E. 1914b. The rare Glossy Ibis in
the Audubon sanctuary at Orange Lake,
Florida. Bird Lore 16:484.
Beadle, D., and A. Whittaker. 1985. 1985 Sabah
survey report. Interwader Ann. Rep. 1985:79-
118,155.
Beals, E.W. 1970. Birds of a euphorbia-acacia
woodland in Ethiopia. 1. Habitat and seasonal
changes./ Anim. Ecol. 39:277-297.
Beasley, A. 1978. Behaviour of Hamerkops.
Honeyguide 94:46.
Beavan, R.C. 1868. Notes on various Indian
birds. Ibis (new ser.) 4:370-406.
Beavan, R.C., and B. Survey. 1865. Indian
ornithological notes, chiefly on the migration
of species. Proc. Zool. Soc. Lond. 1865:690-695.
Beaver, D.L., R.G. Osborn, and T.W. Custer.
1980. Nest-site and colony characteristics of
wading birds in selected Atlantic coast
colonies. Wilson Bull. 92:200-220.
Becher, L. 1967. Der heilige Ibisvoegel der
Aegypter in der Antike. Acta Antique 15:377-
385.
Bechtold, I. 1969-1970. Nesting of Black Storks
(Ciconia nigra) in the surroundings of Koszeg.
Aquila 76-77:187.
Becker, JJ. 1985a. Additions to the late
Pleistocene avifauna of Bradenton, Manatee
County, Florida. Fla. Sci. 47:203-204.
Becker, JJ. 1985b. A late Pleistocene
(Wisconsinan) avifauna from West Palm
Beach, Florida. Bull. Br. Orn. Club 105:37-40.
Beddard, F.E. 1884. A contribution to the
anatomy of Scopus umbretta. Proc. Zool. Soc. Lond.
1884:543-553.
Beddard, F.E. 1886. Notes on the convoluted
trachea of a Curassow (Nothocrax urumutum),
and on the syrinx in certain storks. Proc. Zool.
Soc. Lond. 1886:321-325.
Beddard, F.E. 1888. On certain points in the
visceral anatomy of Balaeniceps rex, bearing
upon its affinities. Proc. Zool. Soc. Lond.
1888:284-290.
Beddard, F.E. 1896. A note upon Dissura
episcopus, with remarks upon the classification
of the herodiones. Proc. Zool. Soc. Lond.
1896:231-235.
Beddard, F.E. 1898. The Structure and Classification
of Birds. Longmans, Green, and Company,
London.
Beddard, F.E. 1901. Some notes on the anatomy
and systematic position of the ciconiine genus
Anastomus. Proc. Zool. Soc. Lond. 1901:365-371.
Bede, P. 1926. Notes sur 1'ornithologie du
Maroc. Mem. Soc. Sci. Nat. Maroc. 16:25-150.
Bednorz, J. 1974. The Black Stork, Ciconia nigra
(L.), in Poland. Ochr. Przyr. 39:201-243.
Beebe, C.W. 1902. Preliminary observations on a
subdermal mite occuring among birds of New
York Zoological Park. Science 15:754-755.
Beebe, C.W. 1907-1915. An ornithological
reconnaissance of northeastern Venezuela.
Zoologica (NY) 1:67-114.
Beebe, C.W. 1914. Notes on the ontogeny of the
White Ibis (Guara alba). Zoologica (NY) 1:241-
248.
Beebe, C.W., and M.B. Beebe. 1910. Our Search
for a Wilderness. Holt and Company, New York.
Beehler, B.M., T.K. Pratt, and D.A.
Zimmerman. 1986. Birds of New Guinea.
Princeton University Press, Princeton.
Beesley, J.S.S. 1971. A waterbird colony in
Tanzania. Bull. E. Afr. Nat. Hist. Soc. 5:73.
Beesley, J.S.S. 1976. New distributional data: 7.
Ostrich 47:214-222.
Beetham, B. 1910. The Home-life of the Spoonbill,
the Stork, and Some Herons. Witherby and Co.,
London. 47pp, 32 pis.
Bekle, H. 1982. Sacred Ibis Threskiornis aethiopica
in Southwestern Australia. West. Aust. Nat.
15:13-19.
Belcher, C.F. 1930. The Birds ofNyasaland. Crosby
Lockwood and Son, London.
Belcher, C., and G.D. Smooker. 1934. Birds of
the colony of Trinidad and Tobago. Ibis (Ser.
13) 4:572-595.
Belknap, H.W. 1957. Observations on the White-
faced Ibis (Plegadis chihi) in Louisiana. M.S.
332
Thesis, Louisiana State University, Baton
Rouge, Louisiana.
Bell, H.L. 1963. Distribution of the Jabiru in
south-eastern Australia. Emu 63:201-206.
Bell-Cross, G. 1974. Observations on fish-eating
birds in central Africa. Honeyguide 77:23-31.
Below, T.H. 1979. First report of pellet ejection
in eleven species. Wilson Bull. 91:626-628.
Belser, C.G. 1989. A Scarlet Ibis or hybrid White
Ibis X Scarlet Ibis in South Carolina. Chat
53:90-91.
Belterman, R.H.R., and L.E.M. de Boer. 1984. A
karyological study of 55 species of birds,
including karyotypes of 39 species new to
cytology. Genetica 65:39-82.
Belton, W. 1984. Birds of Rio Grande do Sul,
Brazil. Part 1. Rheidae through Furnariidae.
Bull. Am. Mus. Nat. Hist. 178:369-636.
Beneda, S. 1988. The White Stork (Ciconia
ciconia) in the West Bohemian District, 1979-
1984. Zpr. Muz. Zapadoces. Kraje Prir. 36-
37:65-87.
Benedict, F.G., and E.L. Fox. 1927. The gaseous
metabolism of large wild birds under aviary
conditions. Proc. Am. Phil. Soc. 66:511-534.
Bennett, G.F., and M.A. Peirce. 1988.
Morphological form in the avian
Haemoproteidae and an annotated checklist of
the genus Haemoproteus Kruse, 1890, J. Nat.
Hist. 22:1683-1696.
Bennett, G.F., E.G. Greiner, and A.G. Campbell.
1975. Avian haemoproteidae. 5. The
haemoproteids of the family of
Threskiornithidae. Can.J. Zool. 53:634-638.
Benson, B. 1973. The flightless ibis of Maui
opens a new chapter in isle's natural history.
Honolulu Advertiser 21 May: A—3.
Benson, C.W. 1958. The occurrence of the Black
Stork Ciconia nigra (Linnaeus) as a Palearctic
migrant in Nyasaland. Bull. Br. Orn. Club
78:86-87.
Benson, C.W. 1961. The breeding of the Whale-
headed Stork in northern Rhodesia. N. Rhodesia
J. 4:557-560.
Benson, C.W. 1963. The breeding seasons of
birds in the Rhodesias and Nyasaland. Proc.
Int. Orn. Congr. 13:623-639.
Benson, C.W. 1967a. The birds of Aldabra and
their status. Atoll Res. Bull. 118:63-111.
Benson, C.W. 1967b. Recent records of the
White Stork, C. ciconia, from Zambia (formerly
Northern Rhodesia). Vogelwarte 24:38-40.
Benson, C.W., and M.P.S. Irwin. 1963. Some
comments on the 'Atlas of European Birds'
from the Ethiopian aspect. Ardea 51:212-229.
Benson, C.W., and M.P.S. Irwin. 1967. A
contribution to the ornithology of Zambia.
Zambia Mus. Papers 1:1-139.
Benson, C.W., and MJ. Penny. 1971. The land
birds of Aldabra. Phil. Trans. R. Soc. Lond. B
260:417-527.
Benson, C.W., and C.R.S. Pitman. 1958. Further
breeding records from Northern Rhodesia.
Bull. Br. Orn. Club 78:164-166.
Benson, C.W., and C.R.S. Pitman. 1961. Further
breeding records from Northern Rhodesia (No.
2). Bull. Br. Orn. Club 81:156-163.
Benson, C.W., and C.R.S. Pitman. 1964. Further
breeding records from Northern Rhodesia (No.
4). Bull. Br. Orn. Club 84:54-60.
Benson, C.W., and D.L. Serventy. 1956.
Breeding data from Katema Island, Northern
Rhodesia. Ostrich 27:171-172.
Benson, C.W., and C.M.N. White. 1957. Check
List of the Birds of Northern Rhodesia.
Government Printer, Lusaka.
Benson, C.W., R.K. Brooke, and C J. Vernon.
1964. Bird breeding data for the Rhodesias
and Nyasaland. Occ. Papers Nat. Mus. S.
Rhodesia 27B:30-m.
Benson, C.W., R.K. Brooke. RJ. Dowsett, and

M.P.S. Irwin. 1971. The Birds of Zambia.
Collins, London.
Bent, A.C. 1926. Life Histories of North American
Marsh Birds. Smithsonian Institution, United
States Museum Bulletin 135, Washington,
D.C.
Bent, A.C., and M. Copeland. 1927. Notes on
Florida birds. Auk 44:371-386.
Bent, A.C., and H.K. Job. 1904. Untitled letter
to W. Dutcher. Auk 21:128-130.
Bentzien, M.M. 1983. Endangered and
threatened wildlife and plants: proposed
endangered status for the U.S. breeding
population of the Wood Stork. Federal Register
48:8402-8404.
Bentzien, M.M. 1984. Endangered and
threatened wildlife and plants: U.S. breeding
population of the Wood Stork determined to
be endangered. Federal Register 49:7332-7335.
Bereszynski, A. 1977. Spostrzezenia nad
zachowaniem sie bocianow bialych (Ciconia
ciconia) i bocianow czarnych (Ciconia nigra) w
puszczy bialowieskiej. [Remarks on behaviour
of black storks and white storks in mixed
flock.] Notatki Orn. 18:62-63.
Berg, B. 1930. To Africa With the Migratory Birds.
G.P. Putman, USA.
Berg, B. 1940. Abu Markub. Dietrich Reimer,
Berlin, 187 pp.
Berger, A. 1910. In Afrikas Wildkammern. Paul
Parey, Berlin. 312pp.
Berlepsch, H.v., and J. Stolzman. 1892. Resultats
des recherches ornithologiques faites au Perou
par M.Jean Kalinowski. Proc. ZooL Soc. Lond.
1892:371-410.
Bernaldo de Quiros, J.L. 1957. Noticia sobre un
nido Ciconia nigra con otros datos. Ardeola
3:229-233.
Berndt, R., and J. Moeller. 1954. Ergebnisse der
Storchforschung im Braunchweiger Land von
1930-1935. Beitr. Naturkd. Niedersachsens 7:101-
109.
Berndt, R., and J. Moeller. 1958.
Bestandsentwicklung des WeiBstorchs (Ciconia
c. ciconia) im Regierungsbezirk Hildesheim von
1907 bis 1953. Vogelring 27:39-47.
Berndt, R., and F. Rehbein. 1961. Ein halbes
Jahrhundert Brutstatistik vom WeiBstorch
(Ciconia c. ciconia) im Kreis Peine. Vogelwarte
21:128-136.
Bernis, F. 1954. Uber Demographic und
Okologie des Weissen Storches in Spanien.
Vogelwarte 17:158-161.
Bernis, F. 1959. La migracion de las ciguenas
espanolas y de las otras ciguenas
'occidentales'. Ardeola 5:9-80.
Bernis, F. 1966. Aves migradores Ibericas. Segun
anillamientos en Europa. vol. 1, pts 1-3.
Publicacion especial de la Sociedad Espanola
de Ornithologia, Madrid. 488pp.
Bernis, F. 1969. Sobre recientes datos de Platalea
leucorodia en Iberia. Ardeola 13:239-241.
Bernis, F. 1969. Sobre recientes datos de Plegadis
falcinellus en Iberia. Ardeola 13:242-243.
Berruti, A. 1980a. Birds of Lake St. Lucia. South.
Birds S:\-6Q.
Berruti, A. 1980b. Status and review of
waterbirds breeding at Lake St. Lucia.
Lammergeyer 28:1-19.
Berruti, A. 1983. The biomass, energy
consumption, and breeding of waterbirds
relative to hydrological conditions at Lake St.
Lucia. Ostrich 54:65-82.
Berruti, A., F. Joubert, M. Skinner, and R.H.
Taylor. 1977. First breeding record of a
Saddle-billed Stork in Natal. Lammergeyer
23:48.
Berthold, P., and E. Nowak. 1988. Anwendung
der Satelliten-telemetrie zur Erforschung des
Vogelzuges. Pp. 181-193 in Elzen, R., K.-L.
Schuchman, and K. Schmidt-Koenig (eds),
Current Topics in Avian Biology, Proc. Int.
Centennial Meeting of the Deutsche
Ornithologen-Gesellschaft. Verlag der
Deutschen O-G, Bonn.
Bertok, L. 1971. Occurrence of the Glossy Ibis
Plegadis falcinellus on the Tyrnau lowland. Slav.
Nar. Muz.. Prir. Vedy. 17:155.
Beruldsen, G.R. 1972. A nest of the Jabiru.
Sunbird 3:40-42.
Beruldsen, G. 1980. A Field Guide to Nests and Eggs
of Australian Birds. Rigby, Sydney. 448pp.
Beteille, G. 1986. [The Spoonbill (Platalea
leucorodia) in the Seine Estuary (1980-1985)].
Cormorant 5:473-479.
Betlem, J. 1982. Frans-Guyana bedreigt de rode
ibis. Vogeljaar 30:96-97.
Betlem, J. 1984. De waarheid over de huidige
situatie van de Rode Ibis op Trinidad.
Vogeljaar 32:1-9.
Betlem, J., and B. de Jong. 1983. De rode ibis
moord in Frans-Guiana. Vogeljaar 31:192-198.
Bezzel, E. 1974. Ornithologishe
Sommerbeobachtungen aus Kleinasien. Anz.
Orn. Ges. Bayern 7:106-120.
Bezzel, E., and B. Wartmann. 1990. New records
of Bald Ibis (Geronticus eremita) in North
Yemen./ Orn. 131:456-495.
Bhutia, U. 1985. Chick-feeding in ibises at
Keoladeo National Park, Bharatpur. J. Bombay
Nat. Hist. Soc. 82:191-192.
Biaggi, V. 1970. Las Aves de Puerto Rico. Univ. de
Puerto Rico, San Juan.
Bierman, W.H. 1959. Observations
ornithologiques au Maroc. Oiseau et R.F.O.
29:4-39,99-128,221-244.
Bigalke, R. 1948. A note on the breeding of the
White-bellied Stork (Sphenorhynchus abdimii) in
the National Zoological Gardens, Pretoria.
Ostrich 19:200-202.
Bildstein, K.L. 1983. Age-related differences in
the flocking and foraging behavior of White
Ibises in a South Carolina salt marsh. Colonial
Waterbirds 6:45-53.
Bildstein, K.L. 1984. Age-related differences in
the foraging behavior of White Ibises and the
question of deferred maturity. Colonial
Waterbirds 7:146-148.
Bildstein, K.L. 1987. Energetic consequences of
sexual size dimorphism in White Ibises
(Eudocimus albus). Auk 104:771-775.
Bildstein, K.L. 1988. Species profile: White Ibis.
Wild Bird August: 14-17.
Bildstein, K.L. 1990a. Status, conservation and
management of the Scarlet Ibis Eudocimus ruber
in the Caroni Swamp, Trinidad, West Indies.
Biol. Conserv. 54:61-78.
Bildstein, K.L. 1990b. The use of zoo collections
in studies of the feeding ecology and
conservation biology of wading birds (Aves:
Ciconiiformes). AAZPA Reg. Conf. Proc.
1990:353-360.
Bildstein, K.L. 1990c. What a difference a day
makes; Hurricane Hugo and the breeding
ecology of coastal South Carolina wading
birds. Colonial Waterbird Society Newsl. 14(2): 17
(abstr.).
Bildstein, K.L., and I.L. Brisbin, Jr. 1989. Lands
for long-term research in conservation biology.
Conserv. Biol. 4:301-308.
Bildstein, K.L., and P.C. Frederick. 1985. Birds
of the Pharoahs. S. Carolina Wildl. 32:6-10.
Bildstein, K.L., R. Christy, and P. Decoursey.
1982a. Size and structure of a South Carolina
salt marsh avian community. Wetlands 2:118-
137.
Bildstein, K.L., R.L. Christy, and P. DeCoursey.
1982b. Energy flow through a South Carolina
salt-marsh shorebird community. Bull. Wader
Study Group 34:35-37.
Bildstein, K.L., S.G. McDowell, and I.L.
Brisbin. 1989. Consequences of sexual
dimorphism in sand fiddler crabs, Uca
333
pugilator: differential vulnerability to avian
predators. Anim. Behav. 37:133-139.
Bildstein, K., W. Post, P. Frederick, and J.
Johnston. 1990a. The importance of freshwater
wetlands in the breeding ecology of coastal
White Ibises (Eudocimus albus): a lesson for
Scarlet Ibis conservation. Pp. 57-63 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds) The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Spoecial Publication 11,
Slimbridge, Glos, UK.
Bildstein, K.L., W. Post, J.Johnston, and P.
Frederick. 1990b. Freshwater wetlands,
rainfall, and the breeding ecology of White
Ibises in coastal South Carolina. Wilson Bull.
102:84-98.
Billing, A.E. 1977. The first occurrence of the
Yellow-billed Spoonbill (Platalea favipes) in
New Zealand. Notornis 24:192.
Binford, L.C. 1989. A Distributional Survey of the
Birds of the Mexican State ofOaxaca.
Ornithological Monograph 43, American
Ornithologists Union, Washington, DC, USA.
Bingham, C.T. 1876. Anastomus oscitans. Stray
Binsbergen, N. 1962. Faglar pa Texel. Var
Fagelvarld 2l:p\s 4 and 5, pp. 144 and 145.
Bishop, K.D. 1984. A Preliminary Report on the
Reserves of South-east Irian Jaya. Interwader,
Kuala Lumpar.
Biswas, B. 1963. Comments on Ripley's 'A
synopsis of the birds of India and Pakistan'. J.
Bombay Nat. Hist. Soc. 60:679-689.
Biswas, B. 1974. Zoological results of the Daily
Mail Himalayan Expedition 1954: notes on
some birds of Eastern Nepal, y. Bombay Nat.
Hist. Soc. 71:456-495.
Blaauw, F.E. 1893. Letter to the editor of The
Ibis. Ibis 1893:465-466.
Blaauw, F.E. 1900. Notes on the zoological
garden of Berlin. Proc. ZooL Soc. Lond.
1900:299-306.
Blaauw, F.E. 1917. Some notes on the Black-
faced Ibis (Theristicus melanopis). Avic. Mag.
8:146-148.
Blacklock, G.W., and R.D. Slacke. 1978. The
Texas colonial waterbird census, 1973-1976.
Proc. Colonial Waterbird Group 1978:99-104.
Blacklock, G.W., D.R. Blankenship, S. Kennedy,
K.A. King, R.T. Paul, R.D. Slack, J.C. Smith,
and R.C. Telfair, III. 1978. Texas Colonial
Waterbird Census, 1973-1976. F.A. Report Series
no. 15. Texas Parks and Wildlife Department,
Austin, Texas, USA. 109pp.
Blackwelder, E. 1907. Research in China
Expedition of 1903-1904 report on zoology.
Carnegie Inst. Wash. Publ. 54, Birds 1(2):483-
506.
Blackwell, A. 1990. The White Stork: a capsule
overview of its status in the wild and the role of
captive reared and bred birds in
reintroduction. Avic. Mag. 96:105-114.
Blake, E.R. 1953. Birds of Mexico, a Guide for Field
Identification. University of Chicago Press,
Chicago.
Blake, E.R. 1977. Manual of Neotropical Birds, vol.
I. University of Chicago Press, Chicago. 674
pp.
Blaker, D. 1967. Spoonbills in the Western Cape.
Blaker, D., and J.M. Winterbottom. 1968. Bird
counts at Strandfontein Sewage Works and
Riet Vlei, Cape Town. Ostrich 39:94-104.
Blakers, M., S.J.J.F. Davies, and P.N. Reilly.
1984. Atlas of Australian Birds. Royal Australian
Ornithological Society, Victoria, Australia.
Blakiston, Capt. 1862. On the ornithology of
northern Japan. Ibis 4:244.
Blancou, L. 1939. Le Balaeniceps rex en Afrique
 Equatorial Francaise. Oiseau Rev. Fr. Orn.
9:480-485.
Blancou, L. 1960. Notes sur des oiseaux de
1'Afrique Centrale. Oiseau Rev. Fr. Orn. 30:276-
278.
Blanford, W.T. 1870. Observations on the Geology
and Zoology of Abyssinia. London.
Blanford, W.T. 1898. The Fauna of British India.
Birds— Vol. IV. Taylor and Francis, London.
500pp.
Blanford, W.T. 1900. The secretary on additions
to the menagerie. Proc. Zool. Soc. Lond.
1900:771-775.
Blaszkiewitz, B., and R. Reinhard. 1985.
Bemerkungen zur Haltung des
Schuhschnabels im Berliner Zoo. Gefiederte Welt
109:36-37.
Blencowe, E J. 1962. Abdim's Storks breeding on
cliffs./ E. Afr. Nat. Hist. Soc. Natl Mus. 24:64.
Blest, A.D. 1961. The concept of'ritualisation'.
Pp. 102-124 in Thorpe, W.H., and O.L.
Zangwill (eds), Current Problems in Animal
Behaviour. Cambridge Univ. Press, London.
Bliese, J.C.W. 1972. White-faced Ibis. Nebr. Bird
Rev. 39:76-77.
Blockey, R. 1939. Experiments with storks. Proc.
Int. Orn. Congr. 9:93-96.
Bloesch, M. 1960. Zweiter Bericht uber den
Einsatz algerischer Storche fur den
Storchansiedlungsversuch in der Schweiz. Orn.
5^.57:214-223.
Bloesch, M. 1980. Drei Jahrzehnte
Schweizerischer Storchansiedlungsversuch in
Altreu, 1948-1979. Orn. Beob. 77:167-194.
Bloesch, M. 1982. Sechsergelege beim Weiss-
Storch Ciconia ciconia. Orn. Beob. 79:39-44.
Bloesch, M. 1984. Ablage und Entwicklung
aussergewohnlich grosser Gelege beim
WeiBstorch. Ciconia ciconia. Orn. Beob. 81:277-
283.
Bloesch, M. 1986. Uber 33 Jahre alter
WeiBstorch Ciconia ciconia in Altreu. Orn. Beob.
83:71-76.
Bloesch, M. 1989a. Der Storchansiedlungs-
versuch in Altreu (Schweiz). Pp. 437-444 in
Rheinwald, G., J. Ogden, and H. Schulz
(eds.), Weiss-Storch: Status undSchutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Bloesch, M. 1989b. Gesellschaft zur Forderung des
Storchansiedlungsversuches Altreu Bull.
18(April):4pp.
Bloesch, M. 1990. Gesellschaft zur Forderung des
Storchansiedlungversuches Altreu Bull. 19
(April):4pp.
Bloesch, M., M. Dizerens, and E. Sutter. 1977.
Die Mauser der Schwungfedern beim Weiss-
Storch (Ciconia ciconia). Orn. Beob. 74(4—6):
161-188.
Bloesch, M., W. Boettcher-Streim, and M.
Dizerens. 1987. Ueber die Mauser des
Grossgefieders beim Schwarzstorch Ciconia
nigra. Orn. Beob. 84:301-315.
Blumel, H. 1984. Zur Brutbiologie des
WeiBstorches. Falke 31:128-130.
Blus, L.J., and R.D. Slack. 1979. Organochlorine
residues in six species of estuarine birds from
South Carolina (USA), 1971-1975. Pestic.
Monit.J. 13:56-60.
Blythe, E. 1842. Letter to the zoological society.
Proc. Zool. Soc. Lond. 1842:93-94.
Blythe, E. 1867. The ornithology of India—a
commentary on Dr. Jerdon's 'Birds of India'.
7^3:147-185.
Boano, G. 1981. La Cicogna Bianca in Piemonte.
Presenza, nidificazione e problemi di
conservazione. Riv. Piem St. Nat. 2:59-70.
Block, WJ. 1964. Kinetics of the avian skull./
Morph. 114:1-42.
Bock, WJ. 1990. A special review: Peters'
'Check-list of Birds of the World' and a history
of avian checklists. Auk 197:629-648.
Bock, W., and J. Terborgh. 1957. Breeding of the
Glossy Ibis in Virginia. Bird-Banding 28:98.
Bodenham, R. 1945. Notes on migrants and
resident birds in the North Suez Canal areas
from March 1 to December 5, 1944. Zool. Soc.
Egypt Bull. 7:21-47.
Bodkin, A.E., and C.T. Mattley. 1921. Shooting
notes. Timehri 7:115-125.
Boehm, M. 1930. Ueber den Bau des
jugendlichen Schaedels von Balaeniceps rex
nebst Bemerkungen ueber dessen
systematischen Stellung und ueber das
Gaumenskelet der Vogel. Z. Morphol. Oekol.
Tiere 17:678-718.
Boekel, C. 1980. Birds of Victoria River Downs
Station and of Yarralin, Northern Territory.
Part \.Aust. Bird Watcher 8:171-193.
Boer, L.E.M. de. 1984. New developments in
vertebrate cytotaxonomy VII. A current list of
references on avian karyology. Genetica 65:3-
37.
Boer, L.E.M. de, and J.M. van Brink. 1982.
Cytotaxonomy of the Ciconiiformes (Aves),
with karyotypes of eight species new to
cytology. Cytogenet. Cell Genet. 34:19-34.
Boer, P., and G.R. Monsees. 1967. Avifaunisch
overzicht van het Balgzand en enkele
omliggende terreinen. Limosa 40:188-205.
Boettcher-Streim, W. 1986. Der
Wiederansiedlungsversuch des Weiss-Storchs
in Altreu/Schweiz. Beih. Veroff. Naturschutz
Landschaftspflege Bad.-Wurtt. 43:315-328.
Boettcher-Streim, W., and M. Bloesch. 1989.
Passage migration of the White Stork Ciconia
ciconia in Switzerland, August and September
1988. Orn. Beob. 86:176-177.
Boettcher-Streim, W., and E. Schuz. 1989.
Bericht uber die IV. Internationale
Bestandsaufnahme des WeiBstorchs 1984 und
Vergleich mit 1974 (6. Ubersicht). Pp. 195-
220 in Rheinwald, G., J. Ogden, and H.
Schulz (eds.), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp. Walsrode
1985, Schriftenr. DDA, Braunschweig,
Germany.
Boettcher-Streim, W., and B. Weissert. 1986.
Zur Frage schmutzverstopfter Schnabel beim
jungen Weiss-Storch (Ciconia ciconia). Okol.
Vogel 8:243-246.
Boettcher-Streim, W., J. Kusche, and E. Sutter.
1990. Von Stoerchen und Menschen. Neidhart and
Schoer, Zurich.
Boev, N., and M. Paspaleva-Antonova. 1964.
Beitrag zur Untersuchung des Schwarzen
Storches (Ciconia nigra L.) in Bulgarien. Izv.
Zool. Inst., Sofya 16:5-16.
Bogachik, T.A., and N.A. Graber. 1987.
Breeding of the White Stork (Ciconia ciconia) in
southwestern regions of the Ukraine.
Ornitologiya 22:201-201.
Bohme, F., and U. Zuppke. 1974. Der Bestand
des WeiBstorchs (Ciconia ciconia) im Kreis
Whittenberg (Bezirk Halle) von 1955 Bis 1970.
Beitr. Vogelkde 20:67-104;.
Bokermann, W.C.A., and J.C.C. Guix. 1987.
Reaparecuimento de Guara, Eudocimus ruber no
litoral de Sao Paulo. Anais Encontro Nac.
Anilhadores Aves 2:206-207.
Bokermann, W.C.A., and J.C. Guix. 1990. Novas
obervagoes sobre a ocorrencia do guara,
Eucocimus ruber, no litoral paulista (Aves,
Threskiornithidae). Anais Encontro Nac.
Anilhadores Aves6A2.
Bolster, R.C. 1923. Note on the breeding season
of the Painted Stork (Pseudotantalus
leucocephalus).J. Bombay Nat. Hist. Soc. 29:561-
562.
Bolster, R.C. 1931. Diet and feeding habits of the
African Ibidae. Ostrich 2:18-19.
Bona, F.V. 1975. Etude critique et taxonomique
des Dilepididae Fuhrm., 1907 (Cestoda).
Parasites des Ciconiiformes. Considerations
334
sur la specifite et la speciation. Monit. Zool.
Ital.Monogr. 1:1-750.
Bonaparte, C.L. 1857. Conspectus Generum Avium,
vol. 2. Brill, Lugduni Batavorum, Leiden.
Bond, J. 1961. Birds of the West Indies. Hough ton
Mifflin Company, Boston.
Bond, J. 1974. Birds of the West Indies, 3rd edn.
Collins, London.
Bonde, K. 1981. An Annotated Checklist to the Birds
of Lesotho. Privately published, Herning,
Denmark.
Bonvallot, J., and G. Randrianasolo. 1975.
Presence du Bec-ouvert Anastomus lamelligerus
sur les Hautes-Terres malgaches. Alauda
43:323-324.
Booser, J., and A. Sprunt. 1980. A literature
review and annotated bibliography of the
Great Basin/Rocky Mountain population of
the White-faced Ibis. U.S. Fisheries and
Wildlife Service Report, Portland.
Borden, D. 1950. Wood Ibis seen near Beaufort,
Carteret Co., N.C. Chat 14:57.
Bordignon, L. 1988. Nesting of the Glossy Ibis,
Plegadisfalcinellus, in Piedmont (N.W. Italy),
Riv. ltd. Orn. 58:186-188.
Boreiko, V.Y. 1988. Operation Stork. Pamphlet
produced by Kiev Reklama, Kiev, USSR.
Borowski, S. 1963a. [Reasons of catastrophical
low nesting success of White Stork in
Bialowieza, 1962]. Notatki Orn. 4:27-29.
Borowski, S. 1963b. Wplyn czynnikow
ekologicznych na legi Bociana Bialego Ciconia
ciconia (L.) w Bialowiezy w roku 1960. Przeg.
ZooL 7:60-62.
Bos, L. v.d. 1982. Agonistisch gedrag bij ibissen.
Een exploratief veldonderzoek. [Agonistic
behavior in ibis. An explorative field research.]
Postgraduate essay, University of Amsterdam,
Amsterdam.
Boswall, J. 1986. To the editor. Oriental Bird Club
Bull. 4:27-28.
Boswall, J. 1989. Some birds at three Chinese
zoos (Xi'an, Nanjing and Nanchang). Avic.
Mag. 95:31-36.
Boswall, J. 1971. Notes from coastal Eritrea on
selected species. Bull. Br. Orn. Club 91:81-84.
Boswall, J.H.R., and M.E.W. North. 1967. A
discography of bird sound from the Ethiopian
zoogeographical region. Ibis 109:521-533.
Bouet, G. 1938a. Le problems de la migration
des Cigognes blanches (Ciconia ciconia ciconia
(L.)) de 1'Afrique du Nord. Proc. Int. Orn.
Congr. 8:671-677.
Bouet, G. 1938b. Nouvelles recherches sur les
Cigognes blanches de 1'Afrique du Nord.
Oiseau Rev. Fr. Orn. 8:20-45.
Bouet, G. 1950. La Vie des Cigognes. Braun et Cie,
Paris.
Bouet, G. 1951. Les Cigognes blanches de
1'Afrique du Nord et leurs migrations. Proc. Int.
Orn. Congr. 10:341-343.
Bouet, G. 1954. La Cicogne blanche Ciconia c.
ciconia au Congo Beige. Ann. Mus. Roy. Congo
Beige Tervuren (Belg.) Ser. 8, Sci. Zool. 1:3-8.
Bouet, G. 1955a. Oiseaux de 1'Afrique Tropicale. Pt.
I. Faune de I'Union Francaise, 16. Larose, Paris.
Bouet, G. 1955b. Acquisitions recentes sur la
migration des Cigognes nordafricaines. Proc.
Int. Orn. Congr. 11:637-639.
Bouet, G. 1956a. L'alimentation des jeunes
Cigognes blanches au nid. Role des insectes.
Oiseau Rev. Fr. Orn. 26:59-61.
Bouet, G. 1956b. Une mission ornithologique en
Algerie en 1955. Nouvelles recherches sur les
Cicognes. Oiseau Rev. Fr. Orn. 26:227-240.
Boulart, R. 1879. Sur les sacs aeriens du
Marabout. Bull. Soc. Philomathique 7:129-130.
Bourne, W.R.P. 1959. Notes on autumn
migration in the Middle East. Ibis 101:170-
176.
Bouvier, M. 1980. Observations of the Black
Stork in Hautes-Alpes. Alauda 48:151.
Bowen, W.W. 1926. Catalogue of Sudan Birds, Pt L
Struthionidae to Picidae. Sudan Government
Museum, Khartoum, Sudan.
Boyd, H. 1983. First Western Hemisphere Waterfowl
and Waterbird Symposium. Proceedings of the
International Waterfowl Research Bureau
Symposium held in Edmonton, Canada 25-28
May. Canadian Wildlife Service, Ottawa,
Ontario. .
Bradley, P. 1985. Birds of the Cayman Islands. P.F.
Bradley, British Indies.
Braine, J.W.S. 1974. Openbilled storks breeding
in South West Africa. Ostrich 45:255.
Braithwaite, L.W., and HJ. Frith. 1969.
Waterfowl in an island swamp in New South
Wales. Ill Breeding. CSIRO Wildlife Research
14:65-109.
Bramley, A. 1979. White Storks migrating at
night by utilizing oil flares. British Birds
72(5):229.
Brand, C.J., R.M. Windingstad, L.M. Siegfried,
R.M. Duncan, and R.M. Cook. 1988. Avain
morbidity and mortality from botulism,
aspergillosis, and salmonellosis at Jamaica Bay
Wildlife Refuge, New York, U.S.A. Colonial
Waterbirds 11:284-292.
Bratton, S.P. 1988. Wood Stork use of fresh and salt
water habitats on Cumberland Island National
Seashore, Georgia. Cooperative Studies Unit
Tech. Rep. 50, Institute of Ecology, University
of Georgia, Athens, Georgia, USA. 24pp.
Bratton, S.P. 1990. Boat disturbance of
Ciconiiformes in Georgia estuaries. Colonial
Waterbirds 13:124-128.
Bratton, S.P., and S. Hendricks. 1988 (1990).
Wood Stork nesting, roosting and foraging at
Cumberland Island, Georgia. Oriole 53:17-24.
Bratton, S.P., C. Canalos, and A. Bergeron.
1989. 1988 Surveys for Wood Storks and Least
Terns, Cumberland Island National Seashore.
Cooperative Studies Unit Technical Report
54, Institute of Ecology, University of Georgia.
Brauer, A. 1911. Ueber die Beteutung des
Musculus ambiens fur die Beugung der Zehen
des Voegels. Sitter. Ges. naturf. Fr. Bed., 175-
179.
Bravery, J.A. 1970. The birds of Athertonshire,
Queensland. Emu 70:49-63.
Bray, M.P. 1986. Feeding ecology of White-faced
Ibises in Lahontan Valley, Nevada. Report of the
Nevada Department of Wildlife, Reno,
Nevada.
Bray, M.P., and D.A. Klebenow. 1988. Feeding
ecology of White-faced Ibises in a Great Basin
Valley, U.S.A. Colonial Waterbirds 11:24-31.
Breach, R.M. 1970. A strange battle. Afr. Wildl
24:64-67.
Breeden, S., and B. Breeden. 1982. The drought
of 1979-1980 at the Keoladeo Ghana
Sanctuary, Bharatpur, Rajasthan. J. Bombay
Nat. Hist. Soc. 79:\-37.
Breeden, S., and S. Slater. 1968. Birds of
Australia. Taplinger Publishing Co., New York.
Brehm, A.E. 1854. Zur Fortpflanzungsgeschichte
einiger Voegel Nord-Ost-Afrika's.y. Orn.
l(Extra-Heft):93-105.
Breife, B. 1982. Skuggfiskande svart stork
(Ciconia nigra). Calidris 11:84-86.
Brelsford, W.V. 1942. Migration notes, Chinsale,
Northern Rhodesia. Ostrich 13:38-41.
Brelsford, W.V. 1943a. Field notes on birds of
the Chambeshi and Luangwa areas of
Northern Rhodesia. Ibis 85:158-163.
Brelsford, W.V. 1943b. Field observations,
Northern Province, Northern Rhodesia. Ostrich
14:170-178.
Brelsford, W.V. 1946. Ecological aspects of the
bird life in the Bangweulu area. Ostrich
17:165-171.
Brelsford, W.V. 1947. Notes on the birds of the
Lake Bangweulu area in Northern Rhodesia.
Ibis 89:57-77.
Brethwaite, C.O., and A. Brady. 1982. Wood
Storks at Cape May Point. Cassinia 60:58-59.
Brewster, W. 1883. The Scarlet Ibis in Florida.
Bull. Nuttall Orn. Club 8:185-186.
Brewster, W. 1886. Breeding of the White-faced
Glossy Ibis in Florida. Auk 3:481-482.
Briggs, G.S. 1931. A note on the birds of the
neighbourhood of Mhow.J. Bombay Nat. Hist.
Soc. 35:382-404.
Brimley, H.H. 1941. Unusual North Carolina
records. ;k* 58:106-108.
Brinkmann, M. 1960. Der Weiflstorch (Ciconia
ciconia) im Reg.-Bez. Osnabruck 1958. Veroff.
Naturwiss. Ver. Osnabruck 29:37-54.
Britton, P.L. 1967. Migration across the
Mediterranean observed by radar. Ibis
109:445.
Britton, P.L. 1970a. Birds of the Balovale
District of Zambia. 0^nVA41:145-190.
Britton, P.L. 1970b. Some non-passerine bird
weights from East Africa. Bull. Br. Orn. Club
90:142-144.
Britton, P.L. 1971. Must the Kisumu Heronry be
condemned a second time? Africana 4:20-22,
32-33.
Britton, P.L. 1978. Seasonality, density and
diversity of birds of a papyrus swamp in
western Kenya. Ibis 120:450-466.
Britton, P.L. (ed.). 1980. Birds of East Africa. East
Africa Natural History Society, Nairobi.
Broadley, D.G. 1973. Predation on birds by
reptiles and amphibians in South Eastern
Africa. Honey guide 76:19-21.
Broadley, D.G. 1974. Predation by birds on
reptiles and amphibians in south-eastern
Africa. Honey guide 78:11-19.
Brockman, HJ., and CJ. Barnard. 1979.
Kleptoparasitism in birds. Anim. Behav.
27:487-514.
Brodkorb, P. 1963. Catalogue of fossil birds. Part
1 (Archaeopterygiformes through
Ardeiformes). Bull. Fla. State Mus. Biol. Sci.
7:179-293.
Brodkorb, P. 1972. New discoveries of Pliocene
birds in Florida (abstract). Proc. Int. Orn.
Congr. 15:634.
Broekhuysen, GJ. 1942. The White Stork
problem. Ostrich 13:10-16.
Broekhuysen, GJ. 1957. Breeding of African
Spoonbill Platalea alba (Scop.) in the South
Western Cape. Ostrich 28:236.
Broekhuysen, G. 1965. Nesting of the White
Stork (Ciconia ciconia (L.)) in South Africa.
Vogelwarte 23:5-11.
Broekhuysen, G. 1967. Bird migration in the
most southern part of the African continent.
Vogelwarte 24:6-\6.
Broekhuysen, GJ. 1971. White Storks breeding
in the Bredasdorp district, most southern part
of the wintering quarters. Vogelwarte 26:164-
169.
Broekhuysen, G. 1973. European storks settle in
South Africa. Afr. Wildl. 27:124-126.
Broekhuysen, G J. 1974. The behavior of White
Storks (Ciconia ciconia) hatched in South Africa.
Vogelwarte 27:166-185.
Broekhuysen, G., and D. Uys. 1966. Ueber das
Brueten des Weiss-Storchs in Suedafrika in
derBrutzeit 1965-66. Vogelwarte 23:235.
Broinowski, GJ. 1890. The Birds of Australia.
Stuart, Melbourne.
Brooke, R.K. 1959. African Spoonbill Platalea
alba breeding in the South Western Cape.
Ostrich 30:88.
Brooke, R.K. 1966. A preliminary list of the
birds of the Kafue National Park. Puku 4:57-
85.
Brooke, R.K. 1969. The calls of Abdim's Storks.
Ostrich 40:22.
Brooke, R.K. 1984. South African Red Data Book.
South African National Scientific Programmes
335
Report 97, December 1984, Cape Town, South
Africa.
Brooke, R.K. in press. The Rare and Vulnerable
Birds of South Africa.
Brooke, R.K., and A.M.B. Masterson. 1971.
Further records of African Marabous breeding
south of the Zambesi. Honeyguide 67:11-14.
Brooke, R.K., B.G. Donnelly, and M.P.S. Irwin.
1982. Comments on cave paintings supposedly
depicting Whale-headed Stork in the Matopos,
Zimbabwe. Honeyguide 110:4-7.
Brookes, J. 1833. On the remarkable formation of
the trachea in the Egyptian Tantalus. Linn. Soc.
Lond. Trans. 16:499-503.
Brooks, D. 1991. Some notes on the
Ciconiiformes in the Paraguan Chaco. Spec.
Group Storks, Ibises, Spoonbills Newsletter 4(1):4-
5.
Brooks, D.J., M.I. Evans, R.P. Martins, and
R.F. Porter. 1987. The status of birds in North
Yemen-Bald Ibis. Sandgrouse 9:12.
Brosset, A. 1956. Evolution actuelle de 1'avifaune
au Maroc oriental. Bull. Soc. Sci. Nat. Phys.
Maroc 36:299-306.
Brosset, A. 1957. Etude de quelques associations
en Afrique du Nord. Alauda 25:122-132.
Brosset, A. 1960. La vie sociale des oiseaux dans
une foret equatoriale de Gabon. Biol. Gabonica
5:29-69.
Brosset, A. 1961. Ecologie des oiseaux du Maroc
oriental. Trav. Inst. Sci. Cherifien Ser. Zool. 22.
Brosset, A. 1974. La nidification des oiseaux
enfore gabonaise. Architecture, situation des
nids et predation. Terre Vie 28:570-610.
Brosset, A., and J. Dragesco. 1967. Oiseaux
collectes et observes dans le Haut-Ivindo. Biol.
Gabonica 3:59-88.
Brosset, A., and C. Erard. 1976. Premiere
descriptions de la nidification de quatre
especes en foret Gabonaise. Alauda 44:205-
235.
Brosset, A., and J.-J. Petter. 1966. Dynamique
des populations d'oiseaux au Maroc oriental.
Bull. Soc. Sci. Nat. Phys. Maroc. 46:400-405.
Brouwer, G.A. 1964. Some data on the status of
the Spoonbill, Platalea leucorodia L., in Europe,
especially in the Netherlands. Zool. Meded.
(Leiden) 39:481-521.
Brouwer, K. 1991. Waldrapp Ibis Conservation
Meeting, June 1991. Spec. Group Storks, Ibises,
Spoonbills Newsletter 4(2):5.
Brouwer, K., and M.C. Coulter. 1990. Letters
from the co-chairs. Spec. Group Storks, Ibises,
Spoonbills Newsletter 3(1/2): 1-2.
Brouwer, K., and A.M. van Wieringen. 1985.
Notes on the behaviour of two colonies of
Scarlet Ibis Eudocimus ruber in captivity.
Postgraduate essay, University of Amsterdam,
Amsterdam.
Brouwer, K., and A.M. van Wieringen. 1986. De
rode ibis. Een onderzoek naar de ethologie en
ecologie van de rode ibis Eudocimus ruber in het
wild. Postgraduate essay, University of
Amsterdam, Amsterdam.
Brouwer, K., and M. van Wieringen. 1988.
Bibliography of the genus Eudcocimus
(Threskiornithidae, Aves). RIN Contributions to
Research on Management of Natural Resources
1988-1.
Brouwer, K., and M. van Wieringen. 1990a.
Nesting ecology of Scarlet Ibises (Eudocimus
ruber) in a small colony of mixed ciconiiform
birds in the Venezuelan llanos. Pp. 16-45 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Brouwer, K., and M. van Wieringen. 1990b.
Bibliography of the genus Eudocimus. Pp. 157-
 191 in Frederick, P.C., L.G. Morales, A.L.
Spaans, and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Brouwer, K., A.L. Spaans, and M. van
Wieringen. 1990. Rode Ibis: symbool van een
goed beschermingsbeleid. Vogels 55:8— 11.
Browder, J.A. 1976. Water, wetlands and Wood
Storks in southwest Florida. Ph.D.
Dissertation, University of Florida,
Gainesville, Florida.
Browder, J.A. 1978. A modeling study of water,
wetlands and Wood Storks. Pp 325-347 in
Sprunt, A., J.C. Ogden, and S. Winkler (eds),
Wading Birds. Research Report 7, National
Audubon Society, New York.
Browder, J.A. 1984. Wood Stork feeding areas in
southwest Florida. Fla. Field Nat. 12:81-96.
Brown, L. 1970. African Birds of Prey. Hough ton
Mifflin, Boston.
Brown, L.H., and P.L. Britton. 1980. The
Breeding Seasons of East African Birds. East
African Natural History Society, Nairobi.
Brown, L.H., and D.E. Pomeroy. 1984. The age
composition of wild populations of birds in
tropical Africa, as demonstrated by plumage
characters, ringing and marking techniques.
Proc. Pan-Afr. Orn. Congr. 5:97-119.
Brown, L.H., and E.K. Urban. 1969. The
breeding biology of the Great White Pelican
Pelecanus onocrotalus roseus at Lake Shala,
Ethiopia, /to 111:199-237.
Brown, L.H., E.K. Urban, and K. Newman.
1982. The Birds of Africa, vol. 1. Academic
Press, New York.
Bruce, R.G. 1979. Wood Storks over Asheville,
North Carolina. C/w/43:19.
Bruce, M.D. 1982. Occurrence of the Lesser
Adjutant Stork Leptoptilos javanicus on Bali,
Indonesia. Bull. Br. Orn. Club 102:39-40.
Brummit, D.W. 1935. Some nesting observations
in the South-East in the summer of 1934-35. S.
Aust. Orn. 1935:49-51.
Brundenell-Bruce, P.G.C. 1975. The Birds of New
Providence and the Bahamas Islands. Collins,
London.
Bruning, D. 1971. Use of canthaxanthin to
maintain the natural colour of captive birds at
Bronx Zoo. Int. Zoo Yb. 11:215-218.
Brunn, B., and A. Sengier. 1970. The Hamlyn
Guide to Birds of Britain and Europe. Hamlyn,
London.
Bryan, A.L. Jr., and M.C. Coulter. 1987a.
Foraging flight characteristics of Wood Storks
in east-central Georgia, U.S.A. Colonial
Waterbirds W:\57-\6\.
Bryan, A.L. Jr., and M.C. Coulter. 1987b.
Foraging site use by Wood Storks of the
Birdsville Colony. Colonial Waterbird Soc.
Motf/rfter 11(3): 17 (abstr.).
Bryan, A.L. Jr., and M.C. Coulter. 1988. Nest
initiation patterns and aggression observed in
the Birdsville Wood Stork Colony, 1987-1988.
Colonial Waterbird Soc. Newsletter 12:25 (abstr.).
Bryan, A.L. Jr., and M.C. Coulter. 1989. Nest
initiation patterns and aggression observed in
the Birdsville Wood Stork colony, 1987-1988.
Pac. Seabird Group Bull. 16(l):22-23 (abstr.).
Bryan, A.L. Jr., and M.C. Coulter. 1990.
Conspecific aggression and nest abandonment
in the Birdsville Wood Stork colony. Colonial
Waterbird Soc. Newsletter 14(2): 18 (abstr.).
Bryant, H. 1861. Notes on some birds observed
in Eastern Florida south of St. Augustine. Proc.
Boston Soc. Nat. Hist. 7:5-21.
Bryant, H.C. 1919. Evidence as to the food of the
Wood Ibis. Condor 21:236-237.
Buckalew, J.H. 1949. The Eastern Glossy Ibis in
Delaware. Auk 66:196-197.
Buckley, F.G. 1978. Colony site selection by
colonial waterbirds in coastal New Jersey
(USA). Proc. Colonial Waterbird Group 1978:17-
26.
Budich, G. 1969. Episoden aus dem
Storchenleben. Falke 16:194-200.
BufTon, G.L.L. d. 1781. Histoire Naturelle des
Oiseaux 15. L'Imprimerie Royale, Paris.
Buil, M.A.M. 1984. Birds of Manggiri. A
preliminary inventory of birds in the Gadja Mada
University's research area 'Manggiri'south of
Yogyakarta, Indonesia. Report 751, Ag. U.
Wageningen, The Netherlands.
Bull,J., and J. Farrand, Jr. 1977. The Audubon
Society Field Guide to North American Birds. Alfred
A. Knopf, New York.
Bult, D. 1983. Birding trip to northern India.
Unpublished report, Putten, The Netherlands.
Bundy, C.F. 1965. A new Floridian: the Scarlet
Ibis. Audubon 67:84-85.
Burger, J. 1978. The pattern and mechanism of
nesting in mixed species heronries. Pp. 45-58
in Sprunt, A., J.C. Ogden and S. Winckler
(eds), Wading Birds. Research Report 7,
National Audubon Society, New York.
Burger, J. 1978. Competition between Cattle
Egrets and native North American herons,
egrets, and ibises. Condor 80:15-23.
Burger, J. 1979. Resource partitioning: nest site
selection in mixed species colonies of herons,
egrets, and ibises. Am. Midi. Natur. 101:191-
210.
Burger, J., and L.M. Miller. 1977. Colony and
nest site selection in White-faced and Glossy
Ibis. Auk 94:664-676.
Burger, J., and J.R. Trout. 1979. Additional data
on body size as a difference related to niche.
Condor 81:305-307.
Burger, J., D. Gladstone, D. Hahn, and L.M.
Miller 1977. Intra- and interspecific
interactions at a mixed species roost of
Ciconiiformes in San Bias, Mexico. Biol. Behav.
2:309-327.
Burgess. 1855. Notes on habits of some Indian
birds. Proc. Zool. Soc. Lond. 1855:70-74.
Burleigh, T.D. 1958. Georgia Birds. University of
Oklahoma Press, Norman, Oklahoma.
Burmeister, H. 1856. Systematische Uebersicht der
Thiere Brasiliens, vol. 2. Reimer, Berlin.
Burton, M., and C.W. Benson. 1961. The
Whale-headed Stork or Shoebill—legend and
hct.N.Rhod. J.4:411-426.
Burton, R.G. 1921. The Adjutant Stork and
other matters./ Bombay Nat. Hist. Soc. 27:633.
Busche, G. 1974. Schwarzstorch (Ciconia nigra)
rastet 10 Toge im anthropogenen Habitat.
[Black Stork roosts for ten days in an urban
habitat]. Orn. Mitt. 26:27.
Bush, A.O., D.B. Pence, and D.J. Forrester.
1973. Tetrameres (Gynaecophila) williamsi sp.
n. (Nematoda: Tetrameridae) from the White
Ibis, Eudocimus albus, with notes on Tetrameres
(Tetrameres) grusi Shumakovich from the
Sandhill Crane, Grus canadensis. J. Parasitol.
59:788-792.
Bush, A.O., and D.J. Forrester. 1976. Helminths
of the White Ibis in Florida. Proc. Helminthol.
Soc. Wash. 43:17-23.
Busk, G. 1870. The secretary on additions to the
menagerie. Proc. Zool. Soc. Lond. 1870:380-384.
Busto, B., and C. Ramo. 1982. Datos
preliminares sobre la reproduccion del
Corocoro Rojo (Eudocimus ruber) en el Llano
Venezolano. Adas del VIII Congreso
Latinoamericano de Zoologia, Merida, pp. 801-805.
Universidad de los Andes, Merida.
Busto, B., and C. Ramo. 1983. La Corocora o
Ibis Escarlata. Natura 73:14-16.
Buthe, A., W.A. Heidmann, B. Peterat, and W.
Ternes. 1989. Schadstoffbelastung des Weiss-
Storchs Ciconia ciconia durch Schwermetalle,
persistent Pestizide und Industriechemikalien.
336
Pp. 415-422 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weiss-Starch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Butler, A.L. 1905. A contribution to the
ornithology of the Egyptian Soudan. Ibis (ser.
8) 19:301-401.
Butler, A.W. 1930a. Roseate Spoonbill in
Florida. Auk 47:75-76.
Butler, A.W. 1930b. Note on the Roseate
Spoonbill in Florida. Auk 47:416.
Butler, E.A. 1881. A tentative catalogue of the
birds of the Deccan and south Mahratta
country. Stray Feathers 9:367-442.
Buxton, C. 1978. The Shoebill story. Black Lechwe
12:6-17.
Buxton, L., J. Slater, and L.H. Brown. 1978. The
breeding behaviour of the Shoebill or Whale-
headed Stork Balaeniceps rex in the Bangweulu
Swamps, Zambia. E. Afr. Wildl.J. 16:201-220.
Buxton, P.A. 1935. Notes on the birds from
northern Nigeria. Ibis (ser. 13) 5:101-110.
Byrd, M.A. 1978. Dispersal and movement of six
North American Ciconiiforms. Pp. 161-185 in
Sprunt, A., J.C. Ogden, and S. Winckler (eds),
Wading Birds. Research Report 7, National
Audubon Society, New York.
Cabot, S. Jr. 1850. Observations upon the recent
appearance in New England of Ibis Guarauna.
Boston Soc. Nat. Hist. Proc. 3:313-314, 332-333.
Calder, D.R. 1959. Wood Ibis, Ibis ibis. Ostrich
30:163-164.
Caldwell, H.R., and J.C. Caldwell. 1931. South
China Birds. H.M. Vanderburgh, Shanghai,
China.
Camacho, M.G. 1983. Notes on aquatic birds of
Nicaragua, Jabiru Jabiru mycteria (Lichtenstein).
Inst. Nicar. Rec. Nat. Ambiente, Managua,
Nicaragua. 14pp.
Camanni, S., and G. Tallone. 1990a.
Nidificazioni della Cicogna Bianca, Ciconia
ciconia in Piemonte Nel 1988 e 1989. Riv. Ital.
Orn. 60:171-176.
Camanni, S., and G. Tallone. 1990b. II projecto
di reintroduzione della Cicogna Bianca
(Ciconia ciconia) in Italia, 1985-1989 (Aves,
Ciconiidae). Riv. Piem. St. Nat. 11:171-183.
Camanni, S., G. Tallone, and R. Piazza. 1989.
L'"Up-down display" nella Cicogna Bianca
(Ciconia ciconia). Contesti ed analisi strutturale.
Mus. Reg. di Science. Nat. Torino 1990:201-203.
Cambi, D. 1982. Ornithological records in
Foggia Province, Italy, marshlands and
Gargano Promontory. Riv. Ital. Orn. 52:137-
153.
Cameron, R.A.D., L. Cornwallis, MJ.L.
Percival, and A.R.E. Sinclair. 1967. The
migration of raptors and storks through the
Near East in autumn. Ibis 109:489-501.
Campbell, A.J. 1892. The breeding haunts of
White Ibis. Victorian Nat. 8:73-74.
Campbell, K.E. Jr. 1979. The non-passerine
Pleistocene avifauna of the Talara Tar Seeps,
northwestern Peru. Life Sci. Contrib. Roy.
Ontario Mus. 118:1-203.
Campbell, K.E. 1986. Comment on the proposed
grant of precedence to Threskiornithidae
Richmond, 1917 (Aves) over Plataleinae
Bonaparte, 1838. Z. N. (S.) 2136. Bull Zool.
Nomencl. 43:10-11.
Campbells, C.W. 1892. On birds collected in
Corea. Ibis 4:244.
Candeias, D., and A. Araujo. 1989. The White
Stork in Portugal. Pp. 19-28 in Rheinwald, G.,
J. Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork
Conservation Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Capanna, E., M.V. Civitelli, and C. Geralico.
1982. The chromosomes of the Hadada Ibis
and comments on the karyotype evolution in
 Threskiornithidae (Aves, Ciconiiformes).
Cytogenet. Cell Genet. 34:35-42.
Capen, D.E. 1977a. Eggshell thickness
variability in the White-faced Ibis. Wilson Bull.
89:99-106.
Capen, D.E. 1977b. The impact of pesticides on
the White-faced Ibis. Ph.D. Dissertation, Utah
State University, Logan, Utah.
Capen, D.E. 1978. Evaluating the impact of
pesticides on the White-faced Ibis in Utah.
Proc. 1977 Colonial Waterbird Group Conf. 1:196-
201.
Capen, D.E., and TJ. Leiker. 1979. DDE
residues in blood and other tissues of White-
faced Ibis. Environ. Pollut. 19:163-171.
Capen, D.E., and J.B. Low. 1974. Recent studies
of the White-faced Ibis in northern Utah. Proc.
Utah Acad. Sci. Arts Lett. 51:198 (abstr.).
Carey, C., H. Rahn and P. Parisi. 1980. Calories,
water, lipid and yolk in avian eggs. Condor
82:335-343.
Carlson, K., and C. Carlson. 1983. Secrets of the
Black Stork. Country Life 4478:1662-1664.
Carpenter, G.D.H. 1946. The relative edibility
and behaviour of some aposematic
grasshoppers. Entomol. Mon. Mag. 82:5-10.
Carrascai, L., J.C. Alonso, and J.A. Alonso.
1990. Aggregation size and foraging behaviour
of White Storks Ciconia ciconia during the
breeding season. Ardea 78:399-404.
Carrick, R. 1959. The food and the feeding
habits of the Straw-necked Ibis Threskiornis
spinicollis (Jameson), and the White Ibis T.
molucca (Cuvier) in Australia. Aust. CSIRO
Wildl. Res. 4:69-92.
Carrick, R. 1962. Breeding, movements, and
conservation of ibises (Threskiornithidae) in
Australia. Aust. CSIRO Wildl. Res. 7:71-88.
Carriker, M.A., Jr. 1910. An annotated list of the
birds of Costa Rica including Cocos Island.
Ann. Carnegie Mm. 6:314-915.
Gary, R. 1973. A Guide to Birds of Southern Portugal.
Uniao Grafica, Lisbon.
Castan, R., and A. Olier. 1959. Le Tantale ibis,
Ibis ibis (Linne), dans le Sud Tunisien et le
Maroc oriental. Alauda 27:148-150.
Castetter, R.C., and H.O. Hill. 1979. Additions
to the birds of the Nevada (USA) test site.
West. Birds W:22\-M3.
Castroviejo, J. 1969. Observaciones de Platalea
leucorodia en Galacia. Ardeola 13:239.
Catesby, M. 1731. The Natural History of Carolina,
Florida and the Bahama Islands. Catesby,
London.
Cave, F.O., andJ.D. MacDonald. 1955. Birds of
the Sudan. Oliver and Boyd, London.
Cawkell, E.M., and R.E. Moreau. 1963. Notes
on birds in The Gambia. Ibis 105:156-178.
Cayley, N.W. 1931. What Bird is That? A Guide to
the Birds of Australia. Angus and Robertson,
Sydney.
Cayley, N.W. 1968. What Bird Is That? 5th edn.
Angus and Robertson, Sydney. 348pp.
Centro Editor de America Latina. 1983a. El
Cueruillo de Canada. Fauna Argentina 53.
Buenos Aires, Argentina.
Centro Editor de America Latina. 1983b. La
Ciguena comun. Fauna Argentina 15. Buenos
Aires, Argentina.
Cerva, F. 1908. A fiatal golyak itatasa a
feszekben. Das Tranken der jungen Storche im
Nest. Aquila 15:311-312.
Chalmers, M.L. 1986. Annotated Checklist of the
Birds of Hong Kong, 4th edn. Hong Kong Bird
Watching Society, Hong Kong. 279pp.
Chamberlain, R.W., R.E. Kissling, D.D. Stamm,
D.B. Nelson, and R.K. Sikes. 1956.
Venezuelan equine encephalomyelitis in wild
birds. Am.J. Hyg. 63:261-273.
Chamberlain, W.D. 1978. The birds of Turtle
Island, S.C.Chat 42:3-7.
Chan, S. 1991. Oriental White Storks wintering
in Hong Kong, 1990/1991. Spec. Group Storks,
Ibises, Spoonbills Newsletter 4(2):3.
Chantrasmi, V. 1979. Wat phai lorn. Holiday
Time in Thailand (January): 18-26.
Chapin, J.P. 1921. A note on the genus
Lampribis in east and central Africa. Ibis (ser.
11)3:609-610.
Chapin, J.P. 1923. The Olive Ibis of Dubus and
its representative on Sao Tome. Am. Mus.
Novit. 84:1-9.
Chapin, J.P. 1932a. The Birds of the Belgian
Congo. Pt. 1. Bull. Am. Mus. Nat. Hist. 65:1-
756.
Chapin, J.P. 1932b. The Birds of the Belgian
Congo, Pt. 3. Bull. Am. Mus. Nat. Hist. 75A:1-
821.
Chapman, F.M. 1903. A new ibis from Mt.
Kenya, British East African. Ibis XXXL:235-
238.
Chapman, E.A. 1969. Gambian observations,
winter 1946-47. Bull. Br. Orn. Club 89:96.
Chapman, F.M. 1912. A new ibis from Mount
Kenya, British East Africa. Bull. Am. Mus. Nat.
Hist. 31:235-238.
Chapman, F.M. 1914. The Roseate Spoonbill.
Bird Lore 16:214-217.
Chapman, F.M. 1917. The distribution of bird-
life in Colombia; a contribution to a biological
survey of South America. Bull. Am. Mus. Nat.
Hist. 36.
Chapman, F.M. 1918. Notes from a traveler in
the tropics. Bird Lore 20:393-397.
Chapman, F.M. 1921. The distribution of bird
life in the Uruebama Valley of Peru. US Nat.
Mus. Bull. 117.
Chapman, F.M. 1926. Distribution of bird-life in
Ecuador. Bull. Amer. Mus. Nat. Hist. 55.
Chappuis, C. 1980. List of sound-recorded
Ethiopian birds. Malimbus 2:1-15, 82-98.
Chasen, F.N. 1935. Hand list of Malaysian birds.
Bull. Raffles Mus. 11.
Chasen, F.N. 1939. The Birds of the Malay
Peninsula, vol. 4. H.F. and G. Witherby,
London.
Chasen, F.N., and C. Boden-Kloss. 1931. Notes
on Malayan birds. Bull. Raffles Mus. 5:80-82.
Chattopadhyay, B. 1988. To home to roost. The
Telegraph, 29 November, 1988.
Chaudhuri, A.B., and K. Chakrabarti. 1973.
Wildlife biology of the Sunderbans forests. A
study of the birds of the Sunderbans with
special reference to the breeding biology of
Openbilled Stork, Little Cormorant, and large
egrets. Set. Cult. 39:8-16.
Chavda, P.B. 1988. Behavioral and ecological
study of the Indian Black Ibis at Junagadh.
M.Ph. Dissertation, Saurashtra University,
Rajkot, India.
Cheesman, R.E., and W.L. Sclater. 1935. On a
collection of birds from north-western
Abyssinia. Ibis (ser. 13) 5:151-191.
Cheke, A.S. 1967. Further notes on land birds
over the Mediterranean and Red Seas. Ibis
109:442-444.
Cheneval, J. 1984. Grallavis edwardsi (Lydekker,
1891), nouveau genre d'oiseau (Ciconiiformes)
du gisement aquitanien de Saint-Gerard-le-
Puy (Allier, France). Bull. Mens. Soc. Linn. Lyon
53:43-60.
Cheng, T.-H. 1955. A Distributional List of Chinese
Birds. I. Non-Passeriformes. Academia Sinica,
Peking.
Cheng, T.-H. (ed.). 1964. China1's Economic Fauna:
Birds. US Dept. Commerce, Joint Publ. Res.
Serv., Washington, DC, USA. 946 pp.
Cheng, T.-H. 1973. The Birth of the Tsing-lin
Mountain Range. Zoological Institute Ac. Sinica,
Peking, China.
Cheng, T.-H. 1976. A Distributional List of Chinese
Birds. Peking Science Press, Peking. 1218pp.
Cheng, T.-H. 1987. A Synopsis of the Avifauna of
China. Science Press, Beijing. 1222pp.
337
Cheylan, G. 1975. Nesting by the stork in the
13th century Milan. Alauda 42:501.
Chikatsuji, H. 1892. Wild Toki, now eight birds.
Anima 37:5.
Child, G. 1964. Observations of aggressive
competition for food among birds in the
Kariba basin. Ostrich 35:55-57.
Child, G. 1972. A survey of mixed 'heronries' in
the Okavango Delta, Botswana. Ostrich 43:60-
62.
Child, P. 1969. Glossy Ibis in Central Otago.
Notornis 16:62.
Child, P. 1973. Far-inland sightings of spoonbill
and godwit. Notornis 20: 1 70-1 7 1 .
Chisholm, A.H. 1934. Bird Wonders of Australia.
Angus and Robertson Limited, Sydney.
Chisholm, A.H. 1958. The diaries of S.W.
Jackson. Emu 58:101-123.
Chisholm, A.H., and V. Serventy. \913.John
Gould's Birds of Australia. Lansdowne Press,
Melbourne.
Choi, G.J. 1966. A visit to sea-bird breeding
places on West Sea. Korean Nature 1966:15-17.
Chozas, P. 1983. Estudio general sobre la
dinamica de la poblacion de la Ciguena
Blanca, Ciconia c. ciconia (L.), en Espana.
Thesis Doctoral, Editorial de la Universidad
Complutense. Servicio de Reprografia,
Madrid.
Chozas, P. 1984. Situacion de la problacion de la
Ciguena blanca, Ciconia ciconia, en Espana
segun los ultimos censos nacionales (1979 y
1981). Bol. Estac. Cent. Ecol. 13:29-48.
Chozas, P. 1985. Mortality rates of Spanish
White Storks (Ciconia ciconia). Ardeola 32:1 19-
123.
Chozas, P. 1986a. Fortpflantzungs- Parameter
des Weiflstorchs (Ciconia ciconia) in
verschiedenen Zonen Spaniens. Beth. Veroff.
Naturschutz Landschaftspflege Bad.-Wurtt 43:221-
234.
Chozas, P. 1986b. Status und Verbreitung des
WeiBstorchs in Spanien 1981 und fruher. Beth.
Veroff. Naturschutz Landschaftspflege Bad.-Wurtt
43:181-187.
Chozas, P. and E. Lazaro. 1988. Causas de
mortalidad de la Ciguena blanca Iberica segun
las recuperaciones de anillsa. Simposio sobre
Ciguena Ibericas, Guadelajaea, Abril 16-19,
1987.
Chozas, P., M. Fernandez-Cruz, and F. Lazaro.
1989. 1984 national census of the White Stork
in Spain. Pp. 29-40 in Rheinwald, G.J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork
Conservation Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Christy, R.L., K.L. Bildstein, and P. DeCoursey.
1981. A preliminary analysis of energy flow in
a South Carolina salt marsh: wading birds.
Colonial Waterbirds4:96-\Q3.
Chubb, C. 1916. Birds of British Guiana, vol. I.
Bernard Quaritch, London.
Chung, J.R. 1986. First report of North Korean
endangered birds [in Japanese] . Newton Graphic
sci,mag.6:122-129
Cieslak, M. 1988. [Nest sites of Black Stork in
Lasy Janowskie, Prov. Tarnobzeg]. Notatki
Orn. 29(3-^):227-231.
Cintra, R. 1986. Nidificacao e crescimento do
filhote de curicaca, Harpiprion caerulescens
(Aves:Threskiornithidae) no Pantanal
Matogrossense. Resumos XIII Congresso Brasileiro
de Zoologica. Cuiba-SBZ.
Clancey, P. A. 1964a. The Birds of Natal and
Zululand. Oliver and Boyd, London.
Clancey, P. A. 1964b. Storks breeding in
Zululand. A correction. Ostrich 35:67.
Clancey, P. A. 1965. A catalogue of birds of the
South African sub-region. (Part I: Families
Spheniscidae— Burhinidae). Durban Mus. Novit.
7:210-304.
Clancey, P.A. (ed.). 1980. South African
Ornithological Society Checklist of Southern African
Birds, 2nd edn. Southern African
Ornithological Society.
Clark, B. 1979. The Bald Ibis. Lanarius 11:4-8.
Clark, C.T. 1982. The Jabiru in the United
States. Birding 14:8-9.
Clark, C.R. 1955. The Sacred Ibis. Bull Metrop.
Mus. Art I3(5)'.m-m.
Clark, E.S. 1978a. Factors affecting the initiation
and success of nesting in an east-central
Florida Wood Stork colony. Proc. Colonial
Waterbird Group 2:178-188.
Clark, E.S. 1978b. Wood Storks using White
Pelicans as beaters. Fla. Fid Nat. 6:45-46.
Clark, E.'S. 1979. The attentiveness and time
budget of a pair of nesting Wood Storks. Proc.
Colonial Waterbird Group 3:204-215.
Clark, E.S. 1980. Attentiveness and time budget
of a pair of nesting Wood Storks. M.S. Thesis,
University Central Florida, Orlando, Florida.
Clark, E.S. 1981. Juvenile Peregrine Falcon
swoops on Roseate Spoonbills. Wilson Bull.
93:548.
Clark, G.A., Jr. 1969. Spread-wing postures in
Pelecaniformes, Ciconiiformes, and
Falconiformes. Auk 86:136-139.
Clark, R.A. 1979a. Seasonal levels of body fat,
protein, ash, and moisture in the Sacred Ibis.
0*^50:129-133.
Clark, R.A. 1979b. DDT contamination of the
Sacred Ibis. Ostrich 50:134-138.
Clark, R.A. 1979c. The food of the Sacred Ibis
Threskiornis aethiopicus at Pretoria, Transvaal.
Clark, R.A., and Clark, A. 1979. Daily and
seasonal movements of the Sacred Ibis at
Pretoria, Transvaal. Ostrich 50:94-103.
Clarke, S.R. 1904. Field-notes of birds obtained
or observed at Bloemfontein, O.R.C., and at
Ingogo, Natal, in 1901 and 1902. Ibis (ser. 8)
4:519-541.
demons, C., and Malinowski, J. 1980. Roseate
Spoonbill in Comanche County, Oklahoma.
Bull. Okla. Ornithol. Soc. 13(3):20-21.
Clinard, T.N. 1979. Wood Storks in western
Virginia. Raven 50:53.
Clinning, C.F., and R.A.C. Jensen. 1977.
Additions to the bird checklist of Etosha
National Park. Madoqua 10:143-148.
Cmak, J. 1989. [The 'Lugi' ornithological reserve
in the region of the Holy Cross Mountains].
Chronmy Przyr. Ojczysta 45:2 1-27.
Coates, BJ. 1985. The Birds of Papua New Guinea,
vol. 1. Non- Passerines. Dove Publ., Alderley,
Queensland, Australia.
Coe, M.J. 1960. Inflation of the neck pouch of
the Marabou Stork. Nature 188:598.
Coghlan, A. 1966. A day in the Drakensberg in
search of Bald Ibis. Bokmakierie 18:43.
Collar, N.J., and P. Andrew (eds). 1988. Birds to
watch, the ICBP world checklist of threatened birds.
1CBP Tech. Publ. 8.
Collar, NJ., and S.N. Stuart. 1985. Threatened
Birds of Africa and Related Islands. The ICBP/
IUCN Red Data Book, Part 1. ICBP,
Cambridge, and IUCN, Gland, Switzerland.
Collar, N.J., P.D. Round, and D.R. Wells. 1986.
Distribution in Thailand. Forktail: 1:33.
Collins, C.T. 1964. Fossil ibises from the
Rexroad fauna of the Upper Pliocene of
Kansas. Wilson Bull. 76:43-49.
Collman, J.R., and J.P. Croxall. 1967. Spring
migration at the Bosphorus. Ibis 109:359-372.
Colodushko, B.Z. 1958. [On biology of the White
Stork]. Trudy Zapoved. Khoz. Belovezh. Pushta
M/w* 1:1 10-1 19.
Colston, P.R. 1971, Additional non-passerine
bird weights from East Africa. Bull Br. Orn.
C/iiA 91:110-111.
Comer, J. A. 1985. The movement and foraging
behavior of Wood Storks in east-central
Georgia. M.S. Thesis. University of Georgia,
Athens, Georgia. 107pp.
Comer, J.A., M.C. Coulter and A.L. Bryan, Jr.
1987. Overwintering locations of Wood Storks
captured in east-central Georgia. Colonial
Waterbirds W:\62-\66.
Comfort, A. 1962. Survival curves of some birds
in the London Zoo. Ibis 104:115-117.
Condamin, M. 1974. Note sur 1'Ibis Rouge en
Guyane. Unpublished report, Office de la
Recherche Scientifique et Technique Outre-
Mer, Cayenne. 9 pp.
Condamin, M. 1975. Etude ecologique du littoral
guyanais en vue de la creation de reserves
naturelles. Unpublished report, Office de la
Recherche Scientifique et Technique Outre-
Mer, Cayenne.
Condamin, M. 1979. L'Ibis Rouge de Guyane.
Cour. Nat. 61:7-14.
Condon, H.T. 1975. Checklist of the Birds of
Australia, Part J. Non-Passerines. Royal
Australian Ornithological Union, Melbourne.
311pp.
Condy, J.B. 1965. A technique for capturing
Abdim's Storks Sphenorhynchus abdimii
(Lichtenstein). Ostrich 36:121-122.
Condy, J.B. 1966. Arrowhead in Abdim's Stork.
Ostrich 37:231-233.
Cone, W.C., and Hall R.T. 1970. Wood Ibis
found nesting in Okefenokee Refuge. Oriole
35:14.
Congreve, W.M. 1943. Two Andalucian
breeding records. Ibis 85:517-518.
Connor, M.A. 1979. Feeding association between
Little Egret and African Spoonbill. Ostrich
50:118.
Cooke, P., J.H. Grobler, and M.P.S. Irwin. 1978.
Notes on Sacred Ibis breeding and other birds
at a dam on Aisleby Municipal Sewage Farm,
Bulawayo. Honeyguide 96:5-11.
Cooke, W.W. 1913. The distribution and migration of
North American herons and their allies. U.S.
Biological Survey Bull. 45, Washington, DC,
USA.
Cooke, W.W. 1897. The Scarlet Ibis in Colorado.
Auk 34:316.
Cookson, E.J., M.R. Hall, and J. Glover. 1988.
The transport of plasma thyroxine in White
Storks (Ciconia ciconia) and the association of
high levels of plasma transthyretin
(Thyroxine-binding prealbumin) with moult.
J.Endocrinol. 117:75-84.
Cooper, J. 1969. Display in the Wood Ibis.
Honeyguide 59:28-29.
Cooper, J. 1970. Marabou stork washing its food.
Honeyguide 61:31-32.
Cooper, K.H., and K.Z. Edwards. 1969. A
survey of Bald Ibis in Natal. Bokmakierie 21:
4-9.
Cott, H.B. 1947. The edibility of birds. Proc. Zoo/.
Soc. London. 116:371-524.
Cottam, C., and P. Knappen. 1939. Food of
some uncommon North America birds. Auk
56:138-169.
Cottam, P.A. 1957. The pelicaniform characters
of the skeleton of the Shoe-billed Stork,
Balaeniceps rex. Bull. Br. Mus. (Nat. Hist.) Zool.
5:51-71.
Coues, E. 1874. Birds of the Northwest: Handbook of
the ornithology of the region drained by the Missouri
river and its tributaries. U.S. Dept. of the Inter.
Misc. Publ. No. 3, Washington, D.C.
Coues, E. 1879. Letters on Ornithology. No.
24-The Wood Ibis. Chicago Field 11:283.
Coues, E. 1903. Key to North American Birds. Dana
Estes and Co., Boston.
Coulter, M.C. 1986a. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Plant:
1984 annual report. SREL-20/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1986b. Wood Storks of the Birdsville
338
Colony and Swamps of the Savannah River Plant:
1985 annual report. SREL-23/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1986c. Wood Storks of the Savannah
River Plant: Foraging and Breeding Ecology.
Comprehensive Cooling Water Study Final Report.
SREL-24/UC-66e. Savannah River Ecology
Laboratory, Aiken, South Carolina.
Coulter, M.C. 1987a. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Plant:
1986 annual report. SREL-31/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1987b. Foraging and breeding
ecology of Wood Storks in east-central
Georgia. Proc. Southeastern Nongame Endang.
Wildl.Symp. 3:21-27.
Coulter, M.C. 1988. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Plant:
1987 annual report. SREL-33/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1989. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Plant:
1988 annual report. SREL-37/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1990a. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Site: 1989
annual report. SREL-38/UC-66e. Savannah
River Ecology Laboratory, Aiken, South
Carolina.
Coulter, M.C. 1990b. Creation and management
of artificial foraging habitat for Wood Storks.
Pp. 262-267 in Mitchell, R.S, CJ. Sheviak,
and D.J. Leopold (eds). Ecosystem Management:
Rare Species and Significant Habitats. New York
State Museum Bulletin 471.
Coulter, M.C. 1990c. The conservation status of
the Oriental White Stork Ciconia boyciana.
P.209 in Matthews, G.V.T. (ed.), Managing
Waterfowl Populations, Proc. IWRB Symp.,
Astrakhan 1989. IWRB Special Publication
12, Slimbridge, Glos, UK. 230pp.
Coulter, M.C., and A.L. Bryan, Jr. 1988. Field
techniques and methods of data collection used in
studies of wood storks of the Birdsville Colony and
swamps of the Savannah River Plant. SREL-34/
UC-66e. Savannah River Ecology Laboratory,
Aiken, South Carolina.
Coulter, M.C., and K. Brouwer. 1991. Letter
from the Co-chairs. Spec. Group Storks, Ibises,
Spoonbills Newsletter 4(1): 1-2.
Coulter, M.C., and J.A. Rodgers. 1987. The
ecology and conservation of storks. Colonial
Waterbirds 10:129-130.
Coulter, M.C., L. Bryan, J. Comer, L. Huff, S.
Jewell, W. Lee, and L. McAllister. 1985. Notes
on Wood Storks in east-central Georgia. Oriole
50:69-70.
Coulter, M.C., L. Bryan, A. Comer, L. Huff, S.
Jewell, W. Lee, and L. McAllister. 1986.
Wood Storks observed in Aiken, Allendale,
and Barnwell Counties, South Carolina. Chat
50:49.
Coulter, M.C., A.L. Bryan, Jr., H.E. Mackey,
Jr.,J.R.Jensen, and M.H. Hodgson. 1987a.
Mapping of Wood Stork foraging habitat with
satellite data. Colonial Waterbirds 10:178-180.
Coulter, M.C., W.D. McCort, and A.L. Bryan,
Jr. 1987b. Creation of artificial foraging
habitat for Wood Storks. Colonial Waterbirds
10:203-210.
Coulter, M.C., S. Balzano, R.E.Johnson, C.E.
King, and P.W. Shannon. 1989. Conservation
and Captive Management of Storks. Stork Interest
Group, University of Georgia Press, Athens,
Georgia.
Coulter, M.C., A.L. Bryan, Jr., Q. Wang, X.
Hu, and C. Gu, 199la. Observations on
Oriental White Stores and other waterbirds at
 Shengjin Lake, 11-21 January, 1989. Pp. 115-
128 in Coulter, M.C., Q. Wang, and C.S.
Luthin (eds), Biology and Conservation of the
Oriental White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Coulter, M.C., A.L. Bryan, Jr., D.P. Young, Jr.,
K. Brouwer, M.P. Kahl, C.E. King, J.A.
Kushlan, C.S. Luthin, and M. van Wieringen.
1991b. A Bibliography of Storks, Ibises and
Spoonbills. Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Courser, W.D., and J.J. Dinsmore. 1975.
Foraging associates of White Ibis. Auk 92:599-
601.
Courtois, R.P.F. 1912. Catalogue des oiseaux du
musee de Zi-Ka-Wei. Mem. Concerhaut I'Hist.
Nat. Empire Chinois 5(3): 1-98.
Coverdale, M.A.C., J. Hancock, and DJ.
Pearson. 1983. Unusual December-January
breeding at the Garsen heronry, Kenya. Scopus
7:49-50.
Cowles, R.B. 1930. The life history of Scopus
umbretta bannermani C. Grant in Natal, South
Africa. Auk 47:159-176.
Cowles, R.B., W.J. Hamilton, III, and F.
Heppner. 1967. Black pigmentation:
adaptation for concealment or heat
conservation? Science 158:1340-1341.
Cowling, S.J. 1974. Observations on ibis feeding
on the Australian plague locust. Emu 74:256-
257.
Cowling, S.J. 1977. Classification of the wetland
habitats of waterbirds. Aust. Mar. Sci. Bull.
58:15-16.
Cowling, S.J., and K.W. Lowe. 1981. Studies of
ibises in Victoria, Australia, I: records of
breeding since 1955. Emu 81:33-39.
Cox, J.J. 1957. Aiafia Banks—island rookery.
Audubon Mag. 59:12-15.
Cracraft, J. 1973. Systematics and evolution of
the Gruiformes (class Aves). 3. Phylogeny of
the suborder Grus. Bull. Am. Mus. Nat. Hist.
151:1-127.
Craib, C.L. 1979. Black Stork breeding in the
Transvaal. Witwatersrand Bird Club News
106:19.
Cram, E.B., 1927. Bird parasites of the Nematode
suborders Strongylata, Ascaridata, and Spirurata.
U.S. Natl. Mus. Bull. 140, Washington, DC,
USA. 465pp.
Cramp, S. 1966. Studies of less familiar birds:
Black Stork. Brit. Birds 59:147-150.
Cramp, S. (ed.). 1977. Handbook of the Birds of
Europe, the Middle East, and North Africa. The
Birds of the Western Palearctic. Oxford University
Press, Oxford.
Cramp, S., and I.J. Ferguson-Lees. 1963. The
birds of the Danube Delta and their
conservation. British Birds 56:323-339.
Crandall, L.S. 1927. The storks. Bull. New York
Zoo/. Soc. 30:149-168.
Crawford, D.N. 1972. Birds of the Darwin area,
with some records from other parts of
Northern Territory. Emu 72:131-148.
Crawford, R.L. 1987. Change in status of some
birds in southwest Georgia. Oriole 52:62-67.
Crawford, W. 1979. Hand-rearing of the Roseate
Spoonbill. Avic. Mag. 85:23-25.
Crawshaw, R. 1907. The Birds of Tierra del Fuego.
Bernard Quaritch, London.
Creutz, G. 1967. Der Schwarzstorch. Falke
14:159.
Creutz, G. 1970a. Das Vorkommen des
Schwarzstorches (C. nigra L.) in Brandenburg.
VeroffBezirksheimatmuseums Potsdam 18:20-30.
Creutz, G. 1970b. Das Vorkommen des
Schwarzstorches (Ciconia nigra L.) in
Brandenberg. Beitr. Tierwelt 5:21-30.
Creutz, G. 1979a. Ein zweidottriges Ei beim
WeiBstorch (Ciconia ciconia). [A double-yolked
egg of the White Stork (Ciconia ciconia)].
Vogelwarte 30(2): 143- 145.
Creutz, G. 1979b. The sight records of one White
Stork. Ring 100:47-48.
Creutz, G. 1982. Neue Ergebnisse zum Zug des
Schwarzstorches. Falke 29:45-50.
Creutz, G. 1985a. Ungewohnlicher
Horstplatzwechsel bei einem WeiBstorches
(Ciconia c. ciconia). Vogelwarte 33:162— 163.
Creutz, G. 1985b. Die Beeinflussung der
Ansiedlung des WeiBstorches (Ciconia ciconia)
durch sein Zugverhalten. Seevogel (Sonderband:
Festsch. FflK*;6:171-172.
Creutz, G. 1988. Der Weiss-Storch, 2. Erweiterte
Auflage. Neue Brehm-Bucherei, Nr. 375. A.
Ziemsen Verlag, Wittenberg/Lutherstadt.
Creutz, G., and L. Creutz. 1970. Der Bestand
des Schwarzstorches (Ciconia nigra L.) und
seine Entwicklung. Beitr. Vogelkd. 16:36-49.
Critchley, R.A., and JJ.R. Grimsdell. 1970.
Nesting of the Shoebill (Balaeniceps rex Gould)
in the Bangweulu Swamps. Bull. Br. Orn. Club
90:119.
Crivelli, A.J., H. Jerrentrup, and B. Hallmann.
1988. Preliminary results of a complete census
of breeding colonial wading birds in Greece,
Spring 1985-1986. Hellenic Orn. Soc. Newsletter
4:31-32.
Crousaz, G. De. 1968. Plegadis falcinellus and
Falco perigrinus at the Bretolet Pass
Switzerland. Nos Oiseaux 29:30.
Csaba, J. 1981. Data on the nesting of Black
Storks (Ciconia nigra) in County Vas. Aquila
88:27-29.
Cuello, J., and E. Gerzenstein. 1962. Las aves del
Uruguay. Lista sisematica, distribucion y
notas. Comun. Zool. Mus. Hist. Nat. Montevideo
6:1-191.
Cugnasse, J., and J. Goar. 1979. La cigogne
blanche nicheuse en Languedoc-Roussillon.
Cunningham- Van Someren, G.R. 1970.
Animated perches and feeding associations of
birds in the Sudan. Bull. Br. Orn. Club 90:120-
122.
Cunningham-van Someren, G.R. 1976. Bird
notes from Karen. E. Afr. Nat. Hist. Soc. Bull.
1976:134-136.
Currie, G. 1972. Why do white storks breed in
S.A.? Bokmakierie 24:71-73.
Currie, G. 1982. Positioning of rings on storks.
Safring News 1 1 :39-4 1 .
Currie, M.H. 1974. Hadedah Ibis (Bostrychia
hagedash) new distributional data. Ostrich
45:133.
Curry, P.J., and J.A. Sayer. 1979. The
inundation zone of the Niger as an
environment for Palaearctic migrants. Ibis
121:20-40.
Curry-Lindahl, K. 1956. Simmande vit stork
(Ciconia ciconia}. Var Fagelvarld 15:277-278.
Curry-Lindahl, K. 1960. Ecological studies on
mammals, birds, reptiles and amphibians in
the eastern Belgian Congo. Part 2. Ann. Mus.
Roy. Congo Beige 87: 1-1 70.
Curry-Lindahl, K. 1961. Contribution a 1'etude
des vertebres terrestres en Afrique tropicale.
Explor. Pare. Nat. Albert et Kagera, Miss. Curry-
Lindahl 1:1-331 .
Curry-Lindahl, K. 1970. Spread-wing postures
in Pelecaniformes and Ciconiiformes. Auk
87:371-371.
Curry-Lindahl, K. 1975. Man in Madagascar.
Defenders Wildl. 50:164-169.
Curry-Lindahl, K. 1978. Conservation and
management problems of wading birds and
their habitats: a global overview. Pp. 83-97 in
Sprunt, A., J.C. Ogden, and S. Winckler (eds),
Wading Birds. Research report 7, National
Audubon Society, New York.
Custer, T.W., and C.A. Mitchell. 1989.
Organochlorine contaminants in White-faced
339
Ibis eggs in southern Texas. Colonial Waterbirds
12:126-129.
Custer, T.W., and R.G. Osborn. 1977a. The use of
feeding habitat by a colony of herons, egrets, and ibises
near Beaufort, North Carolina. U.S. Fisheries and
Wildlife Service, Patuxent Wildlife Research
Center, Laurel, Maryland.
Custer, T.W., and R.G. Osborn. 1977b. Wading
birds as biological indicators: 1975 colony survey.
U.S. Fisheries and Wildlife Service Special
Scientific Report 206. Washington, DC, USA.
Custer, T.W., and R.G. Osborn.. 1978. Feeding
habitat use by colonially-breeding herons,
egrets, and ibises in North Carolina. Auk
95:733-743.
Custer, T.W., R.G. Osborn, and W.F. Stout.
1980. Distribution, species abundance, and
nesting-site use of Atlantic Coast colonies of
herons and their allies. Auk 97:591-600.
Cuvier, G. 1804. Memoire sur 1'ibis des anciens
Egyptiens. Ann. Mus. D'Hist. Nat. 4:166-185.
Cypert, E. 1971. Scarlet Ibis at Okefenokee
National Wildlife Refuge. Oriole 36:22-23.
Cyrus, D., and N. Robson. 1980. Bird Atlas of
Natal. University of Natal Press,
Pietermaritzburg.
Daggett, F.S. 1903. Wood Ibis in southern
California. Condor 5:\8-\9.
Daguerre, J.B. 1922. Lista de aves coleccionadas
y observadas en Rosas, F.C.S. Hornero 2:259-
271.
Daguerre, J.B. 1934. Colonias de aves en
nidificacion. Hornero 5:397-402.
Dahms, G. 1972. 26 jahriger Weiss-Storch
(Ciconia ciconia) als Durchzugler beobachtet.
Vogelwarte 26:355-356.
Dallinga, J.H., and M. Schoenmakers. 1984.
Population changes of the White Stork (Ciconia
ciconia ciconia) during the period 1850-1975.
Nederlandse Vereniging tot Bescherming van
Vogels, Zeist.
Dallinga, J.H., and M. Schoenmakers. 1985.
Over de oorzaken van de achteruitgang van de
ooievaar in NW-Europa. Limosa 58:127-128.
Dallinga, J.H., and S. Schoenmakers. 1987.
Regional decrease in the number of White
Storks (Ciconia c. ciconia) in relation to food
resources. Colonial Waterbirds 10:167-177.
Dallinga, J.H., and S. Schoenmakers. 1989.
Population changes of the White Stork since
the 1850s in relation to food resources. Pp.
231-262 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Danforth, S.T. 1936. Invasion of Glossy Ibis in
Puerto Rico. Auk 53:323.
Daniell, J.P.S. 1951. A heron or a stork. Zoo Life
6:28.
Danielsen, F., and H. Skov. 1987. Waterbird
survey results from South-East Sumatra.
Oriental Bird Club Bull. 6:8-11.
Danielsen, F., H. Skov, and U. Suwarman. in
press a. Breeding colonies of waterbirds along
the coastline of Jambi Province, Sumatra,
August 1989. Kukila.
Danielsen, F., A. Purwoko, M J. Silvius, H.
Skov, and W.J.M. Verheugt. in press b.
Breeding habitat of Milky Stork Mycteria cinerea
in South Sumatra, Indonesia. Kukila.
Darazsi, J. 1958. Observations on the White
Stork. Aquila 65:339-340.
Darling, F.F. 1960. Wild Life in an African
Territory. Oxford University Press, London.
160pp.
Darwin, C. 1962. The Voyage of the Beagle.
Doubleday, New York.
Dathe, F., and G. Degen. 1974. Zum Storchzug
auf dem Balkan. Beitr. Vogelkde 20:484-485.
Dathe, H. 1970. Der Argala. Falke 17:430.
Dathe, H., and LA. Neufeldt (eds). 1978. Atlas
 der Verbreitung Palaearktischer Vogel. 7 Lieforung.
Akademie-Verlag, Berlin.
David, A., and M.E. Oustalet. 1877. Birds of
China. Masson, Paris, France.
Davies, G.C. 1911. Notes on the birds of the
district of Matatiele, East Griqualand. J. S.
Afr. Orn. Union 7:23-48.
Davies, D.H. 1979. Photographs of New York
state rarities 31. White Ibis. Kingbird 29:74-75.
Davis, W.A., and AJ. Reid. 1974. Victorian
ornithological research group Westernport
report no. 1. Part 2. The birds of the Somers,
Sandy Point, Hastings Districts, Westport
Bay, Victoria. Victorian Nat. 91:264-269.
Davis, W.E., Jr. 1979. Analysis of the
vocalization of four heron and ibis species.
Proc. Colonial Waterbird Group 3:216-224.
Davis, W.E,Jr. 1985. Foraging White-faced
Herons follow Australian White Ibises.
Colonial W^rtofr 8:129-134.
Davis, W.M. 1989. Extraordinary aggregation of
White-faced Ibises at a playa lake in Texas
County, Oklahoma. Bull. Okla. Orn. Soc. 22:27.
Dawley, C. 1954. The birds of Guilford County,
North Carolina. Chat 18:30-39.
Dawn, W. 1959. Cattle Egrets provoke cattle to
move and pick flies off bulls. Auk 76:97-98.
Dawson, C.B. 1917. Some colony birds, part III.
Timehri (]. Roy. Agric. Commerc. Soc. Brit.
Guyana) 4:38-57.
Day, D. 1978. Bald ibis (R82) in the southern
Transvaal. Witwatersrand Bird Club News 103:8.
Dean, GJ. 1962. The Whale-headed Stork. N.
Rhodesia J. 5:86-87.
Dean, G J. 1963. Whale-headed Stork Balaeniceps
rex Gould./ E. Afr. Nat. Hist. Soc. 24:77-78.
Dean, GJ.W. 1964. Stork and egret as predators
of the red locust in the Rukwa Valley outbreak
area. Ostrich 35:95-100.
Decary, R. 1950. La Fauna Malgache. Paris.
Payot.
Dechambre, E. 1936. Le Balaeniceps. Terre Vie
6:45-48.
Dee, T J. 1986. The Endemic Birds of Madagascar.
ICBP, Cambridge. 173pp.
Deegtyariov, V.G., G.P. Larionov, and V.N.
Vinokurov. 1990. [The second record of the
Black-headed Ibis (Threskiornis melanocephalus)
in the USSR]. Ornitologiya 24:148.
Deetjen, H. 1964. Nidification de Geronticus
aupres de Ouarzazate. Alauda 32:306-307.
Deetjen, H. 1968. Notes du Moyen Atlas. Alauda
36:287.
Deignan, H.G. 1933. The Jabiru (Jabiru mycteria)
in western Guatemala. Auk 50:429.
Deignan, H.G. 1945. The birds of northern
Thailand. U.S. Natl. Mus. Bull J86,
Washington, DC, USA.
Deignan, H.G. 1963. Checklist of the birds of
Thailand. U.S. Natl Mus. 226:1-263.
Dekeyser, P.L. 1952. Nidification de la cigogne
d'Abdim au Senegal. Notes Afr. 55:92-93.
Dekeyser, P.L. 1954. Contribution a 1'etude du
peuplement de la Mauritanie oiseaux. Bull.
Inst. Fr. Afr. Noire Ser. A 16:1248-1292.
Dekeyser, P.L. 1962. Morpologie et biologic des
oiseaux de 1'Ouest africain. (Notes
documentaries). V. Ardeiformes. Bull. Inst. Fr.
Afr. NoireSer. A Set. Nat. 24a: 1190-1241.
Dekeyser, P.L. 1978. Avifauna aquicola
continental do Brasil (Ensaio de identificacao).
Rev. Nordestina Biol. 1:173-254.
Dekeyser, P.L., and J.H. Derivot. 1966. Les
Oiseaux de I'Quest Africain. ISAN, Dakar.
Delacour, J. 1923. Notes on the birds of the states
of Guarico and Apure in Venezuela. Ibis (ser.
11)5:136-150.
Delacour, J. 1928. On birds collected during the
third expedition to French Indo-china. Ibis
12:23-51.
Delacour, J. 1929. On birds collected during the
fourth expedition to French Indo-china. Ibis
(ser. 12)5:193-220.
Delacour, J. 1932. Les oiseaux de la Mission franco-
anglo-americaine a Madagascar. Oiseau, pp. 1-96.
Delacour, J. 1947. Birds of Malaysia. Macmillan,
New York.
Delacour, J., and J. Greenway. 1940. Liste des
oiseaux recueillis dans la Province du Haut-
Mekong et la Royaume de Luang-Prabang.
Extr. Oiseau Rev. Fr. Orn. 10(1-2):25-59.
Delacour, J., and P. Jabouille. 1925. On the
birds of Quangtri, Central Annam, Ibis (ser.
12) 1:209-260.
Delacour, J., and P. Jabouille. 1931. Les Oiseaux
de L'Indochine Francaise. Expos. Col. Int., Paris.
Delacour, J., and P. Jabouille. 1940. Liste des
oiseaux de 1'Indochine frangaise. Oiseau Rev.
Fr. Orn. (nouv. ser.) 10:89-220.
Delacour, J., and E. Mayr. 1945. Notes on the
taxonomy of the birds of the Philippines.
Zoologica (NY) 30:105-117.
Delacour, J., and E. Mayr. 1946. Birds of the
Philippines. Macmillan, New York.
Delgoda Mendez, J.M. 1981. Policy alternatives
for mangrove ecosystems: a case study of
Laguna de Tacarigua. Unpublished report,
Cornell University, Ithaca, New York.
Delic, A., and I. Matijevic. 1989. [The
population of the White Stork (C. ciconid) near
Grubisno Polje]. Larus40:1-10.
Dementiev, G.P. 1948. Rukovodstvo k Opredeleniiu
Ptits 1948 U.S.S.R. Nauka, Moscow.
Dementiev, G.P., and N.A. Gladkov. \95l.Ptitsy
Sovetskogo Soyuza [Birds of the Soviet Union], vol.
II. Sovetskaya Nauka, Moscow. [Trans, by
Israel Program for Scientific Translations,
Jerusalem, 1969.]
Dementiev, G.P., and N.A. Gladkov. 1960.
Systema Avium rossicarum [Catalogue
critique des Oiseaux d'U.R.S.S.]. Vol. II.
Oiseau Rev. Fr. Orn. 30:1-247.
Dennison, M.D., and H.A. Robertson. 1979.
Records of birds at Farewell Spit, New
Zealand, 1974-1978. Notornis 26:204-207.
Denny, W. 1847. A few remarks on the
geographical distribution of birds in the West
Indies. Proc. Zool. Soc. Lond. 1847:36-41.
Department of the Interior. 1990. Ten foreign
animals proposed for listing as endangered.
Endang. Species Tech. Bull. 15(2):4.
Desai, J.H. 1971. Feeding ecology and nesting of
Painted Storks Ibis leucocephalus at Delhi Zoo.
Int. Zoo Yb. 11:208-215.
Desai, J.H., R.V. Shah, and G.K. Menon. 1974
[1976]. Diet and food requirements of Painted
Storks at the breeding colony in the Dehli
Zoological Park. Pavo 12:13-23.
Desai, J.H., G.K. Menon, and R.V. Shah. 1977
[1978]. Studies on the reproductive pattern of
the Painted Stork, Ibis leucocephalus, Pennant.
Pavo 15:1-32.
De Santo, T.L., S.G. McDowell, and K.L.
Bildstein. 1990. Plumage and behavioral
development of nesting White Ibises. Wilson
Bull. 102:226-238.
Desenne, P., and T.K. Shimotake. 1986.
Observations on the Scarlet Ibis (Eudocimus
ruber) population in the Turuepano area of
Sucre, Venezuela, January 1986. Unpublished
report, Colorado College Biology Department,
Colorado.
Desenne, P., and T. Shimotake. 1990. Algunos
aspectos de la conducta alimentaria del
Corocoro Rojo (Eudocimus ruber) en un bosque
de manglar y especulaciones sobre su dieta.
Pp. 46-56 in Frederick, P.C., L.G. Morales,
A.L. Spaans, and C.S. Luthin (eds) The Scarlet
Ibis (Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
340
DesFayes, M. 1987. Evidence for the ancient
presencer of Bald Ibis, Geronticus eremita, in
Greece. Bull. Br. Orn. Club 107:93-94.
Dewar, D. 1905. A list of the birds found in and
around Madras.y. Bombay Nat. Hist. Soc.
16:484-498.
Dewar, D.R. 1951. Letters on the Ornithology of
Buenos Aires by W. H. Hudson. Cornell
University Press, Ithaca, New York.
Dharmakumarsinhji, R.S. 1955. Birds of
Saurashtra. Privately Published, Bombay.
Dickerman, R.W., C.M. Bonacorsa, and W.F.
Scherer. 1976. Viremia in young herons and
ibis infected with Venezuelan encephalitis
virus. Am.J. Epidemiol. 104:678-683.
Dickerson, W. 1951. Roseate Spoonbills in Palm
Beach area. Fla. Nat. 24:81.
Dickey, D.R., and AJ. van Rossem. 1938. The
birds of El Salvador. Zool. Ser., Field Mus. Nat.
Hist. 23:1-609.
Dietze, R., and D. Sperling. 1983. New
observations on the Glossy Ibis Plegadis
falcinellus in East Saxony, East Germany. Abh.
Ber. Naturkundemus Goerlitz 57:13-14.
Dilley, W.E. 1955. Fishing tackle for the birds.
Everglades Nat. Hist. 3:45-49.
Din, N.A., and S.K. Eltringham. 1974. Breeding
of the Pink-backed Pelican Pelecanus rufescens in
Rwenzori National Park, Uganda with notes
on a colony of Marabou Storks Leptoptilos
crumeniferus. Ibis 116:477-493.
Dindo, J., and K. Marion. 1989. The 1989
breeding season on the Alabama Gulf Coast.
Alabama Birdlife 36(2): 7-9.
Dingle, E.S. 1941. A late record for the White
Ibis in South Carolina. Wilson Bull. 53:234.
Dinsmore, S., andJJ. Dinsmore. 1986. White-
faced Ibis nesting in Dickinson Co., Iowa. Iowa
Bird Life 56:120-121.
Disher, P. 1969. Unique Victorian nesting
record. Bird Obs. 448:7.
Disney, H.J. de S., and AJ. Marshall. 1956. A
contribution to the breeding biology of the
Weaver Finch Quelea quelea (Linnaeus) in East
Africa. Proc. Zool. Soc. Lond. 127:379-387.
Dittami, J., and V. Haas. 1982. A family of
White Storks Ciconia ciconia at Lake Nakuru:
The first breeding record for Kenya? Scopus
6:70.
Dixon, J. 1930. The Wood Ibis in Yellowstone
National Park. Condor 32:288-289.
Dixon, J.E.W. 1970. Miscellaneous notes on
South West African birds. Madoqua 2:45-47.
Dobbrick, W. 1932. Laute Rufausserungen
unserer Stoerche. Orn. Monatsber. 40:18—19.
Dod, A.D. de. 1978. Aves de la Republica Dominica.
Museo Nac. de Hist. Nat., Santo Domingo.
Doguchi, M., and F. Ushio. 1973. PCB and
organochlorine pesticide residues in a White
Stork. Misc. Rep. Yamashina Inst. Orn. 7:202-
206.
Dolderer, P. 1953. Zur Nahrungsbiologie des
Weissen Storches. Naturwiss. Monats. 61:292-
295.
Dolderer, P. 1956. Kaefer und andere Beutetiere
als Zeugen fur das Jagdrevier des Weissen
Storchs. Naturwiss. Monats. 64:21-27.
Dominguez, L., et al. 1985. Diet data of the
Black Stork, Ciconia nigra, in western Spain.
AlytesRECN 3:51-56.
Dopson, C.W., Jr., and M.N. Hopkins, Jr. 1969.
Extension of the White Ibis breeding range in
Georgia. Oriole 32:46-47.
Dornbusch, M. 1965. Weiss-Storch weschselt im
Juni das Weibchen. Falke 12:64.
Dornbusch, M. 1967a. Erhaltet unsere Storche!
Zerbster Heimatkal. 10:98-99.
Dornbusch, M. 1967b. Nachedlege beim WeiB-
storch. Vogelwarte 24:146-147.
Dornbusch, M. 1982a. Zur Populationsdynamik
des Weiss-Storches (Ciconia ciconia). Wiss. Z.
Univ. Greifswald, Math. Nat. R. 31:4,74.
Dornbusch, M. 1982b. Stoerche! Folk* 29:222-
233.
Dornbusch, M. 1982c. Zur Populationsdynamik
des Weiss-Storches Ciconia ciconia (L.). Ber.
Vogelwarte Hiddensee 3:19-28.
Dornbusch, M. 1984a. Wir Zahlen unsere Weiss-
Storche! Aufruf zur Bestandsaufnahme 1984.
Mitt. AK Weiss-Storch 40:1.
Dornbusch, M. 1984b. Wieviel Storche? Aufruf
zu einer Bestandsaufnahme des Weiss-Storchs
1984. 4. Internationale Weiss-Storch-
Bestandsermittlung. Falhe 31:126-127, 140.
Dornbusch, M. 1985. Vorlaufige Auswertung der
Erfassung 1984. Mitt. AK Weiss-Storch 47:1.
Dornbusch, M. 1986. Der Bestand des Weiss-
Storchs (Ciconia ciconia) 1985 in der DDR
(Zwischenauswertung). Mitt. AK Weiss-Storch
56:1.
Dornbusch, M. 1987. Der Weiss-Storch in der
Deutschen Demokratischen Republik.
Ergebnisse der 4. Internationalen
Bestandsaufnahme des Weiss-Storches 1984.
Falke 34:281-288.
Dornbusch, M. 1989. Bestandsentwicklung und
Schutz des Weiss-Storchs, Ciconia ciconia, in der
Deutschen Demokratischen Republik. Pp. 61-
63 in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weiss-Storch: Status und Schut^ Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Dornbusch, M. 1990a. Die Bestandsentwicklung
des Weiss-Storchs 1985-1987. Falke 37:143-
147.
Dornbusch, M. 1990b. Der Bestand des Weiss-
Storchs (Ciconia ciconia) 1988 in der DDR. Mitt.
AK Weiss-Storch.
Dorst,J. 1961. The Migration of Birds. Hough ton
Mifflin, Boston.
Dorst, J. 1967. South America: a Natural History.
Random House, New York.
Dorst, J., and F. Roux. 1972. Esquisse ecologique
sur 1'avifaune des Monts du Bale, Ethiopie.
Oiseau Rev. Fr. Orn. 42:203-240.
Dos Santos, J.R., Jr. 1961. [Population study of
the White Stork in Portugal.] An. Facul. Cienc.
Porto. 44:5-55.
Dourojeanni, M., R. Hofman, R. Garcia, J.
Malleaux, and A. Trovar. 1968.
Observaciones preliminares para el manejo de
las aquaticas del Largo Junin, Peru. Pevta For.
Peru 2:3-52.
Dover, C., and S. Basil-Edwards. 1921. A note
on the habits of the Commom Pariah Kite
(Milvus govinda) and the Adjutant Stork
(Leptoptilos dubius). J. Bombay Nat. Hist. Soc.
27:633.
Downing, H. 1983. A White Ibis near Sheridan,
Wyoming. West. Birds 14:207-208.
Dowsett, RJ. 1966. The status of four species of
aquatic birds in Zambia as suggested by
ringing recoveries. Puku 4:129-131.
Dowsett, RJ. 1969. Ringed Sacred Ibis
Threskiornis aethiopica recovered in Zambia.
Puku 5:59-63.
Dowsett, RJ., and A. de Vos. 1965. The ecology
and numbers of aquatic birds on the Kafue
Flats, Zambia. Ann. Rep. Wildfowl Trust 1963-4
16:67-73.
Draffan, R.D.W. et al. 1983. Emu 83:207-234.
Dragesco, J. 1960. Notes biologique sur quelques
oiseaux d'Afrique equatoriale. Alauda 28:81-
92.
Dragesco, J. 196la. Monographies des Oiseaux
du Bane d'Arguin: La Spatule blanche
(Platalea leucorodia). Set. Nat. 44:7pp.
Dragesco, J. 1961b. Observations ethologiques
sur les oiseaux du Bane d'Arguin. Alauda
29:81-98.
Drahokoupil, L. 1970. A stork pecked to death.
Ziva 18:11.
Dresser, H.E. 1866. Notes on the birds of
southern Texas, part II. Ibis 2:23-46.
Dresser, H.E. 1871-1881. A History of the Birds of
Europe, vol. 6. Taylor and Francis, London.
Drost, R. 1932. Weiflstorch auf dem Zug an der
Omankuste, Arabien. Vogelzug 3:39.
Drouilly, P. 1968. Clave de Giconiiformes de
Chile. Noticiario Mensual 13:3-8.
Dubief, J. 1952. Les Cigognes et le Sahara. Bull.
Liaison Saharienne 11:2-10.
Dubois, G., and J.K. Macko. 1972. Contribution
a 1'etude des Strigeidae la Rue, 1926
(Trematoda: Strigeidae) de Cuba. Ann.
Parasitol. Hum. Comp. 47:51-75.
Dubois, P. 1979. Additional data on some species
observed in Morocco. Alauda 47:43-45.
Dubois, P., and L. Duhautois. 1977. Notes sur
1'ornithologie Marocaine. Alauda 45:285-291.
Dubs, B. 1982. Die Voegel des suedlichen Mato
Grosso. Verlag Verbandsdruckerei-Betadruck
AG, Bern, Switzerland.
Dubs, B. 1988. Beobachtungen zur
Fortpflanzungsbiologie des Stirnbandsibis,
Harpiprioncaerulescens.J. Orn. 129:363-365.
Dubus, M. le chevalier. 1838. Note sur 1'ibis
olivacea. Rev. Zoologique 1838:141-142.
Due, L.D. 1984. Existing status of wetland and
wetland birds conservation in Vietnam.
Unpublished report, University of Hanoi.
Due, L.D., and D. Thuy. 1986. The status of rare
and precious bird species (Ciconiiformes and
Pelecaniformes) in three colonies: Vinh
Thanh, Tan Khanh, Tan Hung; Minh Hai
Province, Vietnam. Unpublished report.
Specialist Group on Storks, Ibises and
Spoonbills, c/o M.C. Coulter, Chocorua, New
Hampshire 03817, USA.
Due, L.D. and P. Tran. undated. The Status of
Kampuchean Wetlands. Unpublished report.
Specialist Group on Storks, Ibises and
Spoonbills, c/o M.C. Coulter, Chocorua, New
Hampshire 03817, USA.
Duckworth, F. 1974. The Whale-headed Stork in
Ethiopia. Bull. Br. Orn. Club 94:3-4.
Dugand, A. 1947. Aves del Departamento del
Atlantico, Colombia. Caldasia 4:499-648.
Duhart, F., and M. Deschamps. 1964. Notes sur
1'avifaune du delta central Nigerien. Oiseau
Rev.Fr. Orn. 33:1-99.
Duignan, P.J., C. Nuttall, J.K. Kirkwood, and
PJ.S. Olney. 1988. Husbandry, breeding and
post-embryonic growth of Abdim's Stork
Ciconia abdimii hand-reared at London Zoo. Int.
Zoo Yb. 27:245-252.
Dujardin, J. 1984. Situation actuelle de 1'Ibis
Rouge en Guyane francaise. Unpublished
report, Office de la Recherche Scientifique et
Technique Outre-Mer, Cayenne.
Dujardin, J.-L. 1987. Situation de 1'Ibis Rouge
(Eudocimus ruber) en Guyane Francaise et dans
le territoire d'Amapa (Bresil) en 1987.
Unpublished report, Office de la Recherche
Scientifique et Technique Outre-Mer,
Cayenne.
Dujardin, J.-L. 1990. Status and conservation of
the Scarlet Ibis in French Guiana. Pp. 107-114
in Frederick, P.G., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Dujardin, J.-L. 1991. Scarlet Ibis in French
Guiana. Spec. Group Storks, Ibises, Spoonbills
Newsletter 4(2):5.
Dujardin, J.-L., and O. Tostain. 1985.
Distribution et eflectifs d'oiseaux d'eau sur le
littoral Guyanas. Alauda 53:287-294.
Dumont, P. A. 1931. Summary of bird notes from
Pinellas County, Florida. Auk 48:246-255.
Dumont, P.A. 1955. Wood Ibis at Gibson Island.
Ail. Nat. 11:6-10.
Duncan, C.D. and P.G.Johnson. 1977. First
341
breeding record of White-faced Ibis for
Alabama and a new area ofPlegadis sympatry.
Ala. Birdlife 19W:\6.
Dunger, G.T. 1965. Wase Rock-its history,
geology, fauna and climbs. Nigerian Guide
30:148-184.
Dunnet, G.M., S.W. Kotagama, and AJ.
Watson. 1978. Water birds. Expedition Report
to the Ruhuna National Park—1978,
University of Aberdeen/Colombo.
Dunnet, G.M., P. Shaw, and S.W. Kotagama.
1979. Bird numbers at Kumana Villu, 15-17 July,
1979. Expedition Report to the Ruhuna
National Park-1979, University of Aberdeen/
Colombo.
Dunning, J. 1977. Breeding Woolly-necked
Stork? Witwatersrand Bird Club News 97:8.
Dunning, J.B., Jr. 1984. Body weights of 686species
of North American birds. Western Bird Banding
Association Monograph No. 1. Eldon Publ.,
Cave Creek, Arizona, USA.
Dunstan, F.M. 1976. Roseate Spoonbill nesting
in Tampa Bay, Florida. Fla. Field Nat. 4(2):25-
27.
duPont, J.E. 1971. Philippine Birds. Delaware
Mus. Nat. Hist. Monograph Ser. 2. Greenville,
Delaware, USA.
Dupuy, A.R. 1967. La faune menacee de
1'Algerie et sa protection. Bull. Soc. Sci. Nat.
Phys.Maroc 47:329-354.
Dupuy, A.R. 1972. Prospection faunistique en
helicoptere au-dessus du Delta du Sine-
Saloum (Senegal). Notes Africaines 136:119-
121.
Dupuy, A.R. 1976. Donnees nouvelles
concernant la reproduction de quelques
especes aviennes au Senegal. Oiseau Rev. Fr.
Orn. 46:47-62.
Dupuy, A.R. 1982. Reproduction of the Marabou
Stork (Leptoptilos crumeniferus) in Senegal. Oiseau
Rev. Fr. Ornithol. 52:293.
Dupuy, A., and J. Verschuren. 1978. Note sur les
Oiseaux, principalement aquatiques, de la
region du Pare National du Delta du Saloum
(Senegal). Le Gerfaut 68:321-345.
Duquet, M. 1990. Statut de la Cigogne blanche
(Ciconia ciconia) et de la Cicogne noire (Ciconia
nigra) en France. Unpublished Report,
Ministere de I'Environement, France. 61pp.
Dusi, J.L. 1966. The identification characters of
nests, eggs, and nestlings of some herons,
ibises, and anhingas. Alabama Birdlife 14:2-8.
Dusi, J.L., and R.T. Dusi. 1968a. Evidence for
the breeding of the Wood Stork in Alabama,
1968. Alabama Birdlife 16:14-16.
Dusi, J.L., and R.T. Dusi. 1968b. Ecological
factors contributing to nesting failure in a
heron colony. Wils. Bull. 80:458-466.
Dusi, J.L., and R.T. Dusi. 1978. Survey methods
used for wading bird studies in Alabama. Pp.
207-211 in Sprunt, A., J.C. Ogden, and S.
Winckler (eds), Wading Birds. Research report
7. National Audubon Society, New York.
Dusi, J.L., and R.T. Dusi. 1988. Thirty-five-year
summary of the status of heron colony sites in
Alabama. Alabama Birdlife 35(2,3): 1-11.
Dusi, J.L., and R.D. Dusi. 1989. Summary of
band recoveries from wading birds banded in
Alabama, 1953-1985. Alabama Birdlife 36:3-6.
Dusi, J.L., R.T. Dusi, and D.L. Bateman. 1971a.
The use of mist nets and decoys in heron and
ibis banding studies. Inl. Bird-banding News
43:6-7.
Dusi, J.L., R.T. Dusi, D.L. Bateman, C.A.
McDonald, JJ. Stuart, and J.F. Dismukes.
1971b. Ecologic impacts of wading birds on the
aquatic environment. Water Resources Research
Institute Bulletin 5, Auburn Univ. 117pp.
Dusi, R.T. 1965. Sight record of the Scarlet Ibis
for Alabama. Wilson Bull. 77:401.
Dutton, T.P. 1972. First nesting record of
 Openbill Stork for the Republic of South
Africa. Lammergeyer 17:36-39.
Duval, C.T. 1969. Recent ornithological records
from the Southern Province. Puku 5:223-226.
Dymin, V.A., and N.S. Pankin. 1975. Nesting
and migration of storks (Ciconidae) and cranes
(Gruidae) in the Upper Amur Region.
Ornithological Studies in the Soviet Far East (New
Series) 29(132):263-267.
Eames, J. 1991. White-shouldered Ibis
rediscovered. Spec. Group Storks, Ibises, Spoonbills
Newsletter 4(2) :5.
Easton, S. 1975. Nesting colony of Marabou
Storks at Shinyanga, Tanzania. E. Afr. Nat.
Hist. Soc. Bull. OcL/Nov.:! 10-111.
Easton, S. 1976. Further notes on the nesting
colony of Marabou Storks in Shinyanga,
Tanzania. E. Afr. Nat. Hist. Soc. Bull.3.
Eaton, E.H. 1910. Birds of New York. Mem. 12.
New York State Museum, Albany.
Eckert, K. 1971. Fall observation of a White-
faced Ibis. Loon 43:12.
Eckert, K. 1988. A White-faced Ibis on Salt Lake
weekend. Loon 60:137.
Ecological Research Division, Thailand Institute
of Scientific and Technological Research.
1981. Ecological Studies for Conservation of Shore
Birds in Songkhla Lake, vol. 1. Office of the
National Environmental Board, Bangkok.
Ecological Research Division, Thailand Institute
of Scientific and Technological Research.
1982. Ecological Studies for Conservation of Shore
Birds in Songkhla Lake, vol. 2. Conservation
plan. Office of the National Environmental
Board, Bangkok.
Edgar, A.T. 1973. Classified summarized notes.
Notornis 20:346-376.
Edwards, E.P. 1972. A Field Guide to the Birds of
Mexico. E.P. Edwards, Sweetbriar, Virginia.
Edwards, H.M. 1930. Roseate Spoonbills (Ajaia
ajaja) taken in southern Alabama. Auk 47:555.
Edwards, H.M. 1932. White ibis near
Montgomery, Alabama. Auk 49:459.
Edwards, W.H. 1847. A Voyage up the River
Amazon, Including a Residence at Para. Murray,
London.
Eggeling, WJ. 1951. Ringed birds recovered in
Uganda. Uganda J. 15:17-25.
Einarsson, O., J. Durinck, M. Peterz, and W.
Vader. 1989. [First record of Black Stork
Ciconia nigra in Iceland.] Bliki 8:51—52.
Eisenmann, E., E. Mayr, and K.C. Parkes. 1984.
Threskiornithidae Richmond, 1917 (Aves):
application to place on official list of family-
group names in zoology and to give precedence
over Plataleinae Bonaparte, 1838, and other
competing family-group names. Z.N.(S.) 2136.
Bull. Zoo/. Nomencl. 41:240-244.
Elbin, S.B. 1990. Multiple methods of identifying
individual Waldrapp Ibises (Geronticus eremita).
AAZPA Annu. Conf Proc. 1990: 208-215.
Elgood,J.H. 1982. The Birds of Nigeria, an
Annotated Checklist, British Ornithologists3 Union
Checklist No. 4. British Ornithologist's Union,
London, England.
Elgood, J.H., and F.C. Sibley. 1964. The tropical
forest edge avifauna of Ibadan, Nigeria. Ibis
106:221-248.
Elgood, J.H., R.E. Sharland, and P. Ward. 1966.
Palaearctic migrants in Nigeria. Ibis 108:84—
116.
Elgood, J.H., C.H. Fry, and R.J. Dowsett. 1973.
African migrants in Nigeria. Ibis 115:1-45,
375-411.
Eliot, W. 1892. The Wood Ibis. Oologist 9:143-
144.
Ellenberger, P.M. 1951. Notes on some birds of
the Ndanga (Zaka) district of Southern
Rhodesia. Ostrich 22:17-24.
Elliot, D.G. 1877. Review of the Ibidinae, or
subfamily of the ibises. Proc. Zoo/. Soc. Lond.
1877:477-510.
Elliott, C.C.H. 1983. Unusual breeding records
made from a helicopter in Tanzania. Scopus
7:33-36.
Elliott, C.C.H., and M.J.F. Jarvis. 1970.
Fourteenth ringing report. Ostrich 41:1—117.
Ellis, R. 1958. Observations on the Straw-necked
Ibis. Emu 58:312.
Elmalik, A.S.E.E. 1989. Threats to the White
Stork in the Sudan and conservation priorities.
Pp. 331-336 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Elst, D. v.d. 1988. Description de la Cigogne
Noire (Ciconia nigra) et de la Cigogne d'Abdim
(Ciconia abdimii). Aves 25:171-182.
Eltringham, S.K. 1968. An experiment on the
control of Marabou Storks. E. Afr. Wildl.J.
6:147-148.
Elwell, N. 1970. Marabous in winter—by the
hundred. Bokmakierie 22:69-71.
Emerson, K.C., and R.D. Proce. 1969. A new
species of Plegadiphilus (Mallophaga:
Menoponidae) from the Cayenne Ibis. Fla.
Entomol. 52:161-163.
England, M.D. 1974. Black Storks wing-spread
while feeding. British Birds 67:236-237.
Ennion, H.E. 1962. Notes on birds seen in Aden
and the Western Aden Protectorate. Ibis
104:560-562.
Erard, C. 1963. Le baguage des oiseaux en 1963.
Bull. Centre Rech. Migrations Mammiferes Oiseaux
17:3-54.
Erickson, R. 1977. First record of the Knot
Calidris canutus, and other records, from Belize
(British Honduras). Bull. Br. Orn. Club 97:78-
81.
Erlanger, C.F. v. 1905. Beitrage zur Vogelfauna
Nordostafrikas. J. Orn. 53:42-158.
Ern, H. 1975. Der Brutbestand des WeiBstorchs
(Ciconia ciconia) in einem Abschnitt der
Kroatischen Posavina (Jugoslawien). Larus
26-28:103-109.
Ernst, C.H. 1974. White Ibis in southeastern
Pennsylvania. All. Nat. 29:181.
Erwin, R.M. 1983. Feeding habitats of nesting
wading birds: spatial use and social influences.
Auk 100:960-970.
Erwin, R.M., J. Kushlan, C. Luthin, I. Price,
and A. Sprunt, IV. 1984. Conservation of
waterbirds in the Caribbean Basin: a summary
of a panel discussion. Colonial Waterbirds 7:139-
142.
Erz, W. 1973. Zunahme des Schwarzstorches in
Polen und in der BRD. Ber. Dtsch. Sekt. Int. Rat.
Vogelschutz 13:94.
Escott, C.J., and D.A. Holmes. 1980. The
avifauna of Sulawesi, Indonesia: faunistic
notes and additions. Bull. Br. Orn. Club
100:189-194.
Etchecopar, R.D., and F. Hue. 1964. Les Oiseaux
du Nord de I'Afrique. Edition N. Boubee, Paris.
606pp.
Etchecopar, R.D., and F. Hue. 1967. The Birds of
North Africa. Oliver and Boyd, London. 612pp.
Etchecopar, R.D., and Hue, F. 1978. Les Oiseaux
de Chine, de Mongolie et de Coree, Non Passereaux.
Les editions du Pacifique, Papeete, Tahiti.
585pp.
Evans, A.H. 1909. Birds. Macmillan, London.
Evans, A.H. 1922. Cambridge Natural History:
Birds. Macmillan, London.
Evans, G.H. 1904. The Black Stork? (Ciconia
nigra?). J. Bombay Nat. Hist. Soc. 16:159.
Evans, K. 1983. Hand-rearing a Scarlet Ibis
Eudocimus ruber at Padstow Bird Gardens,
Cornwall. Avic. Mag. 89:215-217.
Evans, S.M., M.A. Cantrell, and A. Cram. 1981.
Patterns of arrival and dispersal from a mixed
communal roost of Sacred Ibises and Marabou
Storks. Ostrich 52:230-234.
342
Ewans, M. 1989. Bharatpur Bird Paradise. H.F.
and G. Witherby, London. 144pp.
Ezealor, A.U. 1985. Parasites and diseases of
Abdim's Storks Ciconia abdimii. Malimbus
7:120.
Faust, R. and C. Scherpner. 1968. Ibisse mit
'Warzen'. Vogel-Kosmos 12:414-415.
Fabaian, L., G. Boros, and M. Janisch. 1979. Pp.
253-255 in Death of White Storks (Ciconia
ciconia) in Budapest Zoo due to Fluke Infections.
Akademie Verlag.
Fad, K.W., and K.P. Hall. 1989. Hand-raising
Scarlet Ibis. Pp. 78-80 in Proc. 15th National
Conference of the American Association of Zoo
Keepers, Inc., 1989. American Association of Zoo
Keepers, Syracuse, New York.
Fain, A. 1974. Hyper parasitism of storks by
hypopi of Hypoderidae with description of
Neottialges ciconiarum new species acarina
sarcoptiformes. Bull. Ann. Soc. R. Beige. Entomol.
109:190-194.
Fain, A., and B.R. Laurence. 1974. A guide to
the heteromorphic deutonymphs or hypopi
(Acarina: Hypoderidae) living under the skin
of birds, with the description of Ibisidectes debilis
gen. and sp. nov. from the Scarlet Ibis. J. Nat.
Hist. 8:223-230.
Fain, A., and B.R. Laurence. 1979. Neottialges
(Pelecanectes) platalea sp. nov. and other
hypoderid mites (Acarina, Astigmata,
Hypoderidae) from the Spoonbill, Platalea
leucorodia L.J. Nat. Hist. 13:333-336.
Falla, R.A. 1958. Some records of Australian
birds in New Zealand, 1957. Notornis 8:28-32.
Falla, R.A., R.B. Sibson, and E.G. Turbott.
1966. A Field Guide to the Birds of New Zealand.
Collins, London. 254pp.
Falla, R.A., R.B. Sibson, and E.G. Turbott.
1978. The New Guide to the Birds of New Zealand
and Outlying Islands. Collins, London. 247pp.
Fan, Z., and Y. Song. 1991. Saving a treasure of
the World—Japanese Crested Ibis. Forestry of
China 1991(5):36-37.
Fan, Z., Y. Song, and B. Lu. 1991. Protection of
the Crested Ibis for ten years. Chinese Wildlife
59:3-5.
Faria-Pereira, F. 1986. One Glossy Ibis Plegadis
falcinellus observed at Boquilobo Marsh
(Golega). Cyanopica3:758-759.
Farnell, G., and P.W. Shannon. 1987. The
breeding of Abdim's Storks at the Audubon
Park Zoo. Colonial Waterbirds 10:251-254.
Farrand,Jr.,J. (ed.). 1983. The Audubon Society
Master Guide to Birding. 1. Loons to Sandpipers.
Alfred A. Knopf, New York.
Farrar, E., Jr. 1951. White Ibis and Wood Ibis at
Macon. Oriole 16:17-18.
Fatio, V. 1906. Sur le Waldrapp 'Corvus sylvaticus*
de Gesner. Ibis (ser. 8). 6(21): 139-144.
Faxon, W. 1896. John Abbot's drawings of the
birds of Georgia. Auk 13:204-215.
Feduccia, J.A. 1967. Ciconia maltha and Grus
americana from the Upper Pliocene of Idaho.
Wilson Bull. 79:316-318.
Feduccia, A. 1973. Fossil birds from the Late
Pleistocene Ingleside Fauna, San Patricio
County, Texas. Condor 75:243-244.
Feduccia, A. 1977. The Whalebill is a stork.
Nature 266:719-720.
Feduccia, A. 1980. The Age of Birds. Harvard
Univ. Press, Cambridge, MA, USA. 196 pp.
Feely, J.M. 1964. Heron and stork breeding
colonies in the Luangwa valley. Puku 2:76-77.
Feeny, P.P., R.W. Arnold, and R.S. Bailey. 1968.
Autumn migration in the South Caspian
region. Ibis 110:35-86.
Fei, D. 1986. [The geographical distribution of
storks (Ciconiidae) and their conservation in
our country]. Anim. Mondo 3(1):71-74.
Fei, D. 1988a. [The composition of the birds in
Zharlong and its changes.] Pp. 14—23 in The
Birds of Zharlong. Chinese Press/Publishing
 House of Forestry of China, Zharlong, NE
China.
Fei, D. 1988b. [The population of White Ibises in
Zharlong and their protection.} The
Communication of the Avian Society of China,
Gewi Yiang.
Fei, D. 1989a. [Observations on breeding success
of Oriental White Storks in Qiqihar suburban
district.] Dongwuxue Yanjiu 10:263-270.
Fei, D. 1989b. [An observation and investigation
on the breeding situation of the Oriental White
Storks in Qiqihar suburban District.] ZooL Res.
10(3):263-270.
Fei, D. 1991. The breeding of one pair of
Oriental White Storks in the Outskirts of
Qiqihar, Heilongjiang Province. Pp. 59-64 in
Coulter, M.C., Q. Wang, and C.S. Luthin
(eds.), Biology and Conservation of the Oriental
White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Fei, D., G. Wu, W. Ping, C. Wu, and T. Xu.
1983. The observation on the White Stork in
the suburban district of Qiqihaer Heilongjiang
Province, China. Chinese J. Zool. 5:10-13.
Fei, D., Y. Tswen, and T. Yui-Po. 1991a.
Introduction to the studies and conservation of
storks in China. Pp. 137-142 in Coulter, M.C.,
Q. Wang, and C.S. Luthin (eds.), In Biology
and Conservation of the Oriental White Stork
(Ciconia boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Fei, D., W. Ping, G. Wu, T. Wu, and T. Xiu.
1991b. Observations on the breeding biology
of the Oriental White Stork (Ciconia boyciana)
near Qiqihar, Heilongjiang Province, China.
Pp. 21-30 in Coulter, M.C., Q. Wang, and
C.S. Luthin (eds.), Biology and Conservation of the
Oriental White Stork (Ciconia boyciana).
Savannah River Ecology Laboratory, Aiken,
South Carolina, USA.
Feierabend, J.S., and J.M. Zelazny. 1987. Pp.
43-45 in Mitigating, restoring, and creating
wetlands. Status Report on Our Nation's
Wetlands, Natl. Wildl. Fed., Washington,
D.C.
Feldman, R. 1965. Der Schwarzstorch in
Westfalen. Decheniana 118:25-30.
Fennell, C.M, and B.F. King. 1964. New
occurrences and recent distributional records
of Korean birds. Condor 66:239-246.
Fernandez, M. 1920. Los pichones de nuestra
ciguena. Hornero 2:35-38.
Fernandez Cruz, M. 1969. Continuada presencia
de Platalea leucorodia en Extremadura en 1968.
Ardeola 13:239.
Fernandez Cruz, M. 1974. Nuevas observaciones
de Espatula (Platalea leucorodia) en Cacares.
Notas ornitologicas. Ardeola 20:326-327.
Fernandez Cruz, M. 1989. Ringing of the White
Stork in Spain. Pp. 269-270 in Rheinwald, G.,
J. Ogden, and H. Schulz (eds.), Weiss-Storch:
Status undSchutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Fernandez Campos, E.J., JJ. Fuertes Gamundi,
and J.L. Fernandez Estellier. 1974. Censos de
aves acuaticas de las costas gallegas durante la
temporada 1975-76. Brana 1:29-76.
Ferrer-Lerin, F. 1974. Notes on nest building of
Ciconia ciconia in the provinces .of Lerida,
Huesca, Zaragoza, and Teruel. Inst. Munic.
Cienc. Nat. Misc. Zool. 3:83-87.
Feudge, J.B. 1903. The Wood Ibis near San
Bernardin. Condor 5:79.
Fey, D., and T. Lebret. 1972. Veranderingen in
de avifauna van de Biesbosch in het eerste jaar
na de uitbanning van het getij. Limosa 45:101-
118.
ffrench, R. 1967. The Scarlet Ibis. Biol.J. Nat.
Hist. Soc. Univ. W. Indies 1:30-36.
ffrench, R. 1973. A Guide to the Birds of Trinidad
and Tobago. Livingston Publ., Wynnewood,
Pennsylvania. 470pp.
ffrench, R. 1978. Birds of the Caroni Swamp and
marshes. Living World: 1978:42^-4.
ffrench, R. 1980. A Guide to the Birds of Trinidad
and Tobago. Horrowood Books, Pennsylvania.
ffrench, R. 1985. A new look at our Scarlet Ibis.
Trinidad Nat. 6(5):30-34, 55, 59.
ffrench, R.P., and F. Haverschmidt. 1970. The
Scarlet Ibis in Surinam and Trinidad. Living
Bird 9:147-165.
Fiedler, G. 1988. Studying White Storks (Ciconia
ciconia) in Israel by reading rings from a
distance. Torgos 7(2):5-10.
Fiedler, G., and A. Wissner. 1989. Weiss-Storch-
Unfaelle an Freileitungen und
Abhilfemassnahmen. Pp. 423-424 in
Rheinwald, G., J. Ogden, and H. Schulz
(eds.), Weiss-Storch: Status und Schutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Field, F. 1920. Breeding of the Black-necked
Stork (Xenorhynchus asiaticus) . J . Bombay Nat.
Hist. Soc. 27:171-172.
Field, F. 1922. Rough list and notes on the birds
found breeding in the Gonda District, Oudh.
J. Bombay Nat. Hist. Soc. 28:753-772.
Field, G.D. 1978. Status of Ciconiiformes in
Sierra Leone. Bull. Niger. Orn. Soc. 14:42-46.
Figgins, J.D. 1925. Twice-told tales. Proc.
Colorado Mus. Nat. Hist. 5:23-31.
Finch, D.W. 1973. Spring migration.
Northeastern Maritime Region. Am. Birds
27:748-751.
Fincham, J.E. 1971. Black Storks breeding near
Lalapanzi. Honey guide 65:25-27.
Findholt, S.L. 1984. Status and distribution of
herons, egrets, and ibises and related species in
Wyoming. Colonial Waterbirds 7:55-62.
Findholt, S.L., and K.L. Berner. 1988. Current
status and distribution of the Ciconiiforms
nesting in Wyoming. Great Basin Nat. 48:290—
297.
Finley, W.L. 1908. Reports of field agents:
Report of William L. Finley. Bird Lore 10:291-
295.
Finsch, O. 1904. Notiz ueber die Wollhalsstorche
(Dissoura). Orn. Monatsber. 12:94.
Fischer, D. 1978. Bin Kolonie des Weiss-Storches
(Ciconia ciconia) im elektrifizierten
Bahngelaende zwischen Jerewan und
Etschmiadsin (Armenien). Beitr. Vogelkde
24(1/2): 107.
Fischer, R. 1953. Wo sind die Storche? Wetter
Leben 5:123-125.
Fischer, W. 1970. Der Schuhschnabel. Die Neue
Brehm-Bucherei (nr. 425), Wittenberg/
Lutherstadt, 88pp.
Fischer-Sigwart, H. 1914. Comatibis eremita Linne
oder Eudocimus eremita L. Der Waldrapp. Orn.
Beob. 11:73-77.
Fisher, J., N. Simon, and J. Vincent. 1969. The
Red Book: Wildlife in Danger. Collins, London.
Fitzgerald, Q. 1970. Marabous breeding in the
Gandavarowe area. Honeyguide 63:27.
Fitzharris, T. 1989. Zum Voegeln. Living Bird Q.
8(3): 15-21.
Fjeldsa, J. 1983. Vertebrates of the Junin Area,
Central Peru. Steenstrupia 8:285-298.
Fjeldsa, J. 1985a. Origin, evolution, and status of
the avifauna of Andean wetlands. Pp. 85-1 12
in Buckley, P.A., M.S. Foster, E.S. Morton,
R.S. Ridgley, and F.G. Buckley (eds),
Neotropical Ornithology. Ornithological
Monograph No. 36. American Ornithologists'
Union, Washington D.C.
Fjeldsa, J. 1985b. Classification of waterbird
communities in South-eastern Australia. Emu
85:141-149.
Fjeldsa, J. 1988. Status of birds of steppe habitats
on the Andean Zone of Patagonia. ICBP Tech.
Publ. 7:88-95.
343
Fjeldsa, J., and N. Krabbe. 1990. Birds of the High
Andes. Zool. Museum, Copenhagen and Apollo
Books, Svendborg, Denmark.
Flam, J. 1989. First Kentucky record of the
Roseate Spoonbill. Ky. Warbler 65:96.
Fleming, R.L. Sr., R.L. Fleming Jr., and L.S.
Bangdel. 1979. Birds of Nepal, 2nd edn. Avalok
Publishers, Katmandu.
Fleming, T. 1990. White Storks at Zeak. Oman
Bird News 8:6-7.
Fleming, W.J., J.A. Rodgers, and CJ. Stafford.
1984. Contaminents in Wood Stork eggs and
their effects on reproduction, Florida, 1982.
Colonial Waterbirds 7:88-93.
Fletcher, K.C. 1979. Repair of bilateral
mandibular fractures in a Shoebill Stork
(Balaeniceps rex).J. Zoo. An. Med. 10:69-72.
Flickenger, E.L., and D.L. Meeker. 1972.
Pesticide mortality of young White-faced Ibis
in Texas. Bull. Environ. Contam. Toxicol. 8:115-
168.
Flint, V.E. 1978. Far Eastern Stork Ciconia
boyciana. Pp. 92-93 in Flint, V.E. (ed.), The Red
Data Book of the U.S.S.R., Part 2: Birds. Lesnaya
Promyshlennost Publishers, Moscow.
Flint, V.E., and S.M. Smirenskii. 1977.
[Preliminary Summary Soviet Map concerning
the Protection of Migratory and Endangered
Birds.] P. 251 in Winter, S. (ed.), VII All-Soviet
Ornithological Conference. Zoological Institute of
the Academy of Science of the Ukranian SSR,
Kiev.
Flint, V.E., and S.M. Smirenskii. 1978. New
data on the distribution of the Japanese Crane
and the Far Eastern White Stork (abstr.) VII
USSR National Ornithological Conference 2:251.
Flint, V.E., R.L. Boehme, Y.V. Kostin, and A.A.
Kuznetsov. 1984. A Field Guide to the Birds of the
USSR. Princeton University Press, Princeton,
New Jersey.
Flower, S.S. 1922. The hermit ibis in the Sudan.
7Hr 11:598-599.
Flower, S.S. 1925. Contributions to our
knowledge of the duration of life in vertebrate
animals. —IV. birds. Proc. Zool. Soc. Lond.
1925:1365-1422.
Flower, S.S. 1938. Further notes on the duration
of life in animals —IV. birds. Proc. Zool. Soc.
Lond. (ser.A) 108:195-235.
Forbes, W.A. 1877. On the bursa fabricii in
birds. Proc. Zool. Soc. Lond. 1877:304-318.
Forbush, E.H., & J.B. May. 1955. A Natural
History of American Birds of Eastern and Central
North America. Bramhall House, New York.
Ford, E.B. 1943. Wood Ibis at Wepener. Ostrich
14:183-184.
Ford, J.R. 1957a. Additional notes on the bird
life of Leonora, Western Australia. Emu 57:21-
24.
Ford, J.R. 1957b. Swan River observations. Emu
57:352-354.
Ford, J.R., and P.S. Stone. 1957. Birds of the
Kellerberrin/Kwolyin District, Western
Australia. Emu 57:9-21.
Forrester, D J. 1980. Hematozoa and
Mallophaga ffom the White Ibis Eudocimus
albus in Florida U.S.A.J. Parasitol. 66:58.
Forrester, D.J., E.G. Greiner, G.F. Bennett, and
M.K. Kigaye. 1977. Avian Haemoproteidae.
7. A review of the Haemoproteids of the family
Ciconiidae (storks) and descriptions of
Haemoproteus brodkorbi sp. nov. and H. peirci sp.
nov. Can.J. Zool. 55:1268-1274.
Fouarge, J. 1987. [Post-nuptial call of the Black
Stork (Ciconia nigra).] Aves 24:152-153.
Fox, D.L. 1962. Carotenoids of the Scarlet Ibis.
Comp. Biochem. Physiol. 5:31^3.
Fox, D.L., and T.S. Hopkins. 1966. Carotenoid
fractionation in the Scarlet Ibis. Comp. Biochem.
Physiol. 19:267-278.
Fradrich, H. 1969. Der Zoo van Tananarive.
Zool. Gart. 37(l/3):41-47.
Francis, E. 1978. White Ibis in Transylvannia
County, N.C.Chat 42:81.
Francke, C., and B. Grefe. 1984. White Ibis
sighted in Saginaw County. Jack-Pine Warbler
62:50.
Frank, C. 1976. Syngamus-trachea as the cause
of helminthiasis in young storks. Angew
Parasitoi. 17:99-100.
Fraser, L. 1843. Notes on species brought to
England by Mr. Bridges. Proc. ZooL Soc. Lond.
1843:108-121.
Fraser, W. 1971. Breeding herons and storks in
Botswana. Ostrich 42:123-127.
Fraser, W. 1982. Feeding associations of
Carmine Bee-eaters with other animals.
Bokmakierie 34:18.
Frederick, P.C. 1982. Mating patterns in White
Ibis. BirdBehav. 4:51 (abstr.).
Frederick, P.C. 1985a. Mating strategies of
White Ibis (Eudocimus albus). Ph.D.
Dissertation, University of North Carolina,
Chapel Hill, North Carolina.
Frederick, P.C. 1985b. Intraspecific food piracy
in White Ibis./ Field Orn. 56:413-414.
Frederick, P.C. 1986a. Conspecific nest takeovers
and egg destruction by White Ibises. Wilson
Bull 98:156-157. •
Frederick, P.C. 1986b. Parental desertion of
nestlings by White Ibis (Eudocimus albus) in
response to muscle biopsy. J. Field Orn.
57:168-170.
Frederick, P.C. 1986c. A self-tripping trap for use
with colonial nesting birds. N. Am. Bird Bander
11:94-95.
Frederick, P.C. 1987a. Extrapair copulations in
the mating system of White Ibis (Eudocimus
albus). Behaviour 100:170-201.
Frederick, P.C. 1987b. Chronic tidally-induced
nest failure in a colony of White Ibis. Condor
89:413-419.
Frederick, P.C. 1987c. Responses of male white
ibises to their mate's extra-pair copulations.
Behav. Ecol. Sociobiol. 21:223-228.
Frederick, P.C. 1990. Hydrological cues
associated with the initiation and
abandonment of nesting by White Ibises
(Eudocimus albus) in the Everglades marshes of
Florida, U.S.A. Pp. 64^70 in Frederick, P.C.,
L.G. Morales, A.L. Spaans, and C.S. Luthin
(eds.), The Scarlet Ibis (Eudocimus ruber):
Status, Conservation., and Recent Research, Proc. 1st
International Scarlet Ibis Conservation
Workshop, Caracas, Venezuela. IWRB
Special Publication 11, Slimbridge, Glos, UK.
Frederick, P.C., and M.W. Collopy. 1988.
Reproductive ecology of wading birds in relation to
water conditions in the Florida Everglades.
Technical Report No. 30. Florida Cooperative
Fish and Wildlife Research Unit, Department
of Wildlife and Range Science, University of
Florida, Gainsville, Florida.
Frederick, P.C., and M.W. Collopy. 1989a.
Nesting success of five ciconiiform species in
relation to water conditions in the Florida
Everglades. Auk 106:625-634.
Frederick, P.C., and M.W. Collopy. 1989b. The
role of predation in determining reproductive
success of colonially nesting wading birds in
the Florida Everglades. Condor 91:860-867.
Frederick, P.C., and M.A. Shields. 1986.
Suspected intraspecific egg dumping in the
White Ibis (Eudocimus albus). Wilson Bull.
98:476-478.
Frederick, P.C., L.G. Morales, A.L. Spaans, and
C.S. Luthin (eds). 1990. El Corocoro Rojo
(Eudocimus ruber): Situacion Actual,
Conservacion, e Investigaciones Recientes [The
Scarlet Ibis (Eudocimus ruber): Status,
Conservation, and Recent Research], Proc. 1st
International Scarlet Ibis Conservation
Workshop, Caracas, Venezuela. IWRB
Special Publication 11, Slimbridge, Glos, UK.
Freligh, S. 1985. 'Father of the Shoe'. Int. Wildl.
15:14^15.
Friedmann, H. 1930. Birds collected by the Childs
Frick Expedition to Ethiopia and Kenya Colony. Part
I. Non-Passerines. U.S. Natl. Mus. Bull. 153,
Washington, DC, USA. 516pp.
Friedmann, H., and A. Loveridge. 1937. Notes
on the ornithology of tropical East Africa. Bull.
Mus. Comp. Zoo/. Harvard 81:1-413.
Friedmann, H., and F.D. Smith, Jr. 1950. A
contribution to the ornithology of northeastern
Venezuela. Proc. U.S. Nat. Mus. 100:411-538.
Friedmann, H., and F.D. Smith, Jr. 1955. A
further contribution to the ornithology of
north-eastern Venezuela. Proc. U.S. Natl. Mus.
104:463-524.
Friedmann, H., and J.G. Williams. 1969. The
birds of the Sango Bay forests, Buddu County,
Masaka District, Uganda. Contrib. Sci (Los
Ang.) 162:1-48.
Friendly, A. 1973. Search for tomb in Egypt
uncovers vast animal cult. Smithsonian 4:66-73.
Frith, HJ. (ed.). 1976. Reader's Digest Complete
Book of Australian Birds. Reader's Digest
Services, Sydney. 615pp.
Frith, H.J., and S.J.J.F. Davies. 1961. The
breeding seasons of birds in subcoastal
Northern Territory. Emu 61:47-111.
Frome, N.F. 1947. The birds of Delhi and
district./ Bombay Nat. Hist. Soc. 47:277-300.
Frohring, P.C., and R.A. Beck. 1978. First
breeding record of the White Ibis Eudocimus
albus in Virginia. American Birds 31:73-74.
Frohring, P.C., D.P. Voorhees, and J.A.
Kushlan. 1988. History of wading bird
populations in the Everglades: a lesson in the
use of historical information. Colonial Waterbirds
11:328-335.
Fruhstuck, H. 1976. Der Bestand des Weiss-
Storches (Ciconia ciconia L.) im Burgenland im
Jahre 1976. [The status of the White Stork
(Ciconia ciconia) in Burgenland in 1976.] Egretta
20(2):61-64.
Fruhstuck, H. 1982. [Population of the white
stork (Ciconia ciconia) in Burgenland, Austria,
during 1979 and 1980.] Egretta 25:33-36.
Fruhstuck, H. 1989. Zur Situation und
Bestandsentwicklung des Weiss-Storchs im
Burgenland. Vogelschutz Osterreich 4:26-28.
Fry, C.H. 1966. Report on wetlands in Nigeria with
reference to wildfowl and waders. IWRB Report.
Fry, C.H. (ed.). 1967. White Storks at Kaduna
and Zaria. Bull. Niger. Orn. Soc. 4:37.
Fry, C.H., S. Keith, and E.K. Urban. 1985.
Evolutionary expositions from The Birds of
Africa: Halcyon song phylogeny; cuckoo host
partitioning; systematics ofAplopelia and
Bostrychia. Pp. 163-180 in Schuchmann, K.-L.
(ed.), African Vertebrates: Systematics, Phylogeny,
and Evolutionary Biology. Zool.
Forschungsinstitut und Mus. A. Koenig,
Bonn.
Fujimaki, Y. 1990. Records of (Ciconia ciconia
boyciana from Hokkaido, Japan. Jap.J. Orn.
37:37-38.
Gabaldon, G., and G. Ulloa. 1979. Ciconiiformes
de Venezuela: clave para su identification y
otras consideraciones utiles en estudios de
malaria aviaria. Bol. Dir. Malariol. Saneamiento
Ambiental 19:84-109.
Gabaldon, G., and G. Ulloa. 1980.
Holoendemicity of malaria: an avian model.
Trans. Roy. Soc. Trop. Med. Hyg. 74:501-507.
Gabrielson, I.N., and S.G. Jewett. 1970. Birds of
the Pacific Northwest. Dover Publications, New
York.
Gadow, H.F. 1887. Remarks on the cloaca and
on the copulatory organs of the Amniota. Phil.
Trans. Roy. Soc. Lond. B. 178:5-37.
Gadow, H.F. 1888. Remarks on the numbers and
on the phylogenetic development of the
344
remiges of birds. Proc. Zool. Soc. Lond.
1888:655-667.
Gadow, H., and E. Selenka. 1891. Vogel. Vol. I.
Anatomischer Theil. In Bronn's Klassen und
Ordnungendes Thier-Reichs. Leipzig. 1008pp.
Galarza, A. 1986. Migracion de la espatula
(Platalea leucorodia (Linn.)) por la peninsula
Iberica. Ardeola 33:195-201.
Galdikas, M.F., and B. King. 1989. Lesser
Adjutant nests in S.W. Kalimantan. Kukila
4:151.
Gallagher, M.D. 1986. Abdim's Stork in Arabia.
Sandgrouse S:\Q7-\ll.
Gallagher, M.D. 1989. The White Stork in
Oman, eastern Arabia. Pp. 327-330 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Gallagher, M., and M.W. Woodcock. 1980. The
Birds of Oman. Quartet Books, New York.
Gandidier, G., and G. Petit. 1932a. New
observations in Madagascar. Alauda 39:37-42.
Gandidier, G., and G. Petit. 1932b. Zoologie de
Madagascar. Paris.
Gangloff, B., and L. Gangloff. 1987. L'elevage de
la Cigogne blanche (Ciconia ciconia) en
captivite. [Raising storks (Ciconia ciconia) in
captivity.] Zool. Garten 57:305-324.
Gangloff, L., B. Gangloff, B. Schmitt, and A.
Schmitt. 1989. Fortpflanzungsprogramm in
Gefangenschaft des WeiBstorchs im
Strassburger Zoo. Pp. 445-460 in Rheinwald,
G., J. Ogden, and H. Schulz (eds), Weiss-
Storch: Status und Schutz, Proc. 1 st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Ganguli, U. 1975. A Guide to the Birds of the Delhi
Area. Indian Council of Agricultural Research,
Delhi.
Ganier, A.F. 1955. Wood Ibis near Tullahoma.
Migrant 26:43.
Garavelli, C. 1991. Riproduzione in captivita dell
ibis eremita (Geronticus eremita). Parco Nat. Viva
Rapp. No. 1.
Garcia, F., and O. Garrido. 1965. Catalogo de las
aves de Cuba. Trabajos de divulgacion, Museo
'Felipe Poey' de la Academia de Ciencias de la
Republica de Cuba, La Habana.
Garcia, L., J.A. Amat, and M. Rodriguez. 1983.
Spoonbills breeding during winter in Spain.
British Birds 76:32-33.
Garcia-Oliva, J., M. Gonzales Nicolas, and JJ.
Aja. 1979. Observacione de Espatulas
anilladas en la marisma de Santona
(Santander). Donana Acta Vert. 6:236-237.
Gardner, L.L. 1925. The adaptive modifications
and the taxonomic value of the tongue in
birds. Proc. U.S. Natl. Mus. 67:1-49.
Gariboldi, A. 1987. Records of Ciconia ciconia and
C. nigra in the province of Pavia (NW Italy) in
1986. Riv. Ital. Orn. 57:130-131.
Garland, I. 1963. Nesting of the Woolly-necked
Stork. Bokmakierie 15(2):15-16.
Garland, I.F. 1981. The history of the Siaya
catchment area dating back to 1945 and the
changes that took place ... as I saw them. Siaya
Proj. News Letter 1:4-7.
Garrido, O.H., and F. Garcia. 1975. Catalogo de
las Aves de Cuba. Academia de Ciencias de
Cuba, Havana. 149pp.
Garrod, A.H. 1873. On the carotid arteries of
birds. Proc. Zool. Soc. Lond. 1873:457-472.
Garrod, A.H. 1873-1874. On certain muscles of
the thigh of birds and on their value in
classification. Proc. Zool. Soc. Lond. 1873:626-
644; 1874:111-123.
Garrod, A.H. 1875a. On the form of the trachea
in certain species of storks and spoonbills. Proc.
Zool. Soc. Lond. 1875:297-301.
Garrod, A.H. 1875b. On the disposition of the
 deep plantar tendons in different birds. Proc.
ZooL Soc. Lond. 1875:339-348.
Garrod, A.H. 1877. Note on an anatomical
peculiarity of certain storks. Proc. ZooL Soc.
Lond. 1877:711-712.
Garrod, A.H. 1878. On the trachae of Tantalus
loculator and of Vanellus cqyennensis. Proc. ZooL
Soc. Lond. 1878:625-631.
Garstone, R. 1973. Yellow-billed spoonbills at
Woodanilling. State Wildl. Advisory News Sew.
[Publ. by Dept Fish. Fauna, Perth, Western
Australia.] 4:39.
Garzon Heydt, J. 1973. Observacione de Platalea
leucorodia en la provincia la Salamanca. Ardeola
19:14.
Gaston, G.R. 1976. Occurrence of four species of
ibis near Dauphin Island, Alabama. Alabama
Birdlife24:\4.
Gaston, G.R., and P.G.Johnson. 1977. Nesting
success and mortality of nestlings in a coastal
Alabama heron/egret colony. Northeast Gulf Set.
1:14-22.
Gates, G.L. 1930. Handbook of the Birds of West
Africa.]. Bales and Sons and Danielson, Ltd,
London, UK.
Gauckler, A., and M. Kraus. 1962. Braune
sichler (Plegadis falcinellus L.) in Franken. Anz.
Orn. Ges. Eayern 6(4):385-388.
Gebauer, A. 1988. Nachgelege beim Weiss-
Storch. Ciconia,Jahresbericht Tierpark Gorlitz
1:22-23.
Gebauer, A. 1989. Zum Problem der
morphologischen Geschlechtsdifferenzierung
beim WeiBstorch (Ciconia ciconia). Ciconia,
Jahresbericht Tierpark Gorlitz 2:18-21.
Gehl, O. 1975. Young stork choked on a ring
snake. Falke 22:13.
Geroudet, P. 1965a. Notes sur les oiseaux du
Maroc. Ardea 33:295-308.
Geroudet, P. 1965b. Du 'Waldrapp' de Gessner
aux ibis chauves du Maroc. Nos Oiseaux
28:129-143.
Geroudet, P. 1978. De nouveau des cigognes en
Suisse trente ans d'une experience de
reintroduction. Nos Oiseaux 34(7):311-318.
Geus, A. 1959. Der Waldrapp in Europa
ausgestorben. Falke 6:133-136.
Geyr von Schweppenburg, H. 1922. Zur Theorie
des Vogelzuges. J. Orn. 70:361-385.
Geyr von Schweppenburg, H. 1936. Storchzug
und Mittelmeer. J. Orn. 84:339-351.
Ghorpade, K.D. 1973. Preliminary notes on the
ornithology of Sandur, Karnataka. J. Bombay
Nat. Hist. Soc. 70:499-531.
Gibson, E. 1880. Ornithological notes from the
neighborhood of Cape San Antonio, Buenos
Ayres. Ibis (ser. 4) 4:153-169.
Gibson, E. 1919. Further ornithological notes
from the neighbourhood of Cape San Antonio,
Province of Buenos Ayres. Part II.
Trochilidae-Plataleidae. Ibis (ser. 11) 1:495-
537.
Gibson, E. 1920. Further ornithological notes
from the neighbourhood of Cape San Antonio,
Province of Buenos Ayres. Part III.
Phoenicopteridae-Rheidae. Ibis (ser. 11) 2:1-
97.
Gibson-Hill, C.A. 1949. An annotated checklist of the
birds of Malaya. Bull. Raffles Mus. 20,
Singapore.
Giebel, G. 1873. Balaeniceps rex. Z. Gesam.
Naturwiss. 41:350.
Gill, H.B. 1970. Birds of Innisfail and
Hinterland. Emu 70:105-116.
Gillespie, P., and L. Arnold. 1982. Pathology
report. Dodo J.Jersey Wildl. Preserv. Trust 19:81-
94.
Gilliard, E.T. 1958. Living Birds of the World.
Doubleday and Co., Garden City, New Jersey.
Girard, G.T. and W.K. Taylor. 1979.
Reproductive parameters for nine avian
species at Moore Creek, Merritt Island
National Wildlife Refuge, Florida. Fla. Sci.
42:94-102.
Giraudoux, P., and E. Schiiz. 1978. Fang von
Weifistorchen auch in Niger. Vogelwarte
29:276-277.
Gizenko, A.I. 1962. [On nidification of the
Glossy Ibises at the Dnieper River.] Orn.
Moscow 6:469.
Glade, A.A. (ed.). 1988. Red list of Chilean
Terrestrial Vertebrates. CONAF, Santiago, Chile.
Gladkov, N.A. 1965. [Research on the fauna of the
Soviet Union (birds).] Moscow University,
Moscow.
Gladstone, D.E. 1979. Promiscuity in
monogamous colonial birds. Amer. Nat.
114:545-557.
Glas, L., and D. Porper. 1986. [Nesting of storks
on the Golan Plateau.] Tzufit (Sunbird) 4:6-15.
Glenister, A.G. 1951. Birds of the Malay Peninsula,
Singapore, and Penang. Oxford University Press,
London.296pp.
Glenny, F.H. 1945. A systematic study of the
main arteries in the region of the heart. Aves
XIII. Ciconiiformes, Pt. I . Am. Midi. Nat.
33:449-454.
Glenny, F.H. 1953. A systematic study of the
main arteries in the region of the heart. Aves
XIII. Ciconiiformes, Pt. 2. Ohio J. Sci. 53:347-
348.
Glenny, F.H. 1955. Modifications of pattern in
the aortic arch system of birds and their
phylogenetic significance. Proc. U.S. Natl. Mus.
104:525-621.
Gliddon, G.R. 1850. Remarks upon the scarcity
of Ibis religiosa in Egypt. Proc. Acad. Nat. Sci.
Phila. 5:83-84.
Glushchenko, Y.N. 1985a. [The problem of
conservation and apealing of Far-eastern Stork
in the Prichankaiskaya lowland.] In Rare and
Endangered Birds of the Far East. Far East
Central Academy of Science of the USSR,
Vladivostok.
Glushchenko. Yu. M. 1985b. Problems of
protection and investigation of Eastern White
Storks in the Khanka Lowland. Pp.l3&-139 in
Rare and Endangered Birds of the Far East. Far
East Central Academy of Science of the USSR,
Vladivostok.
Gochfeld, M. 1973. Observations on new or
unusual birds from Trinidad, West Indies and
comments on the genus Plegadis in Venezuela.
Condor 75:474-478.
Godfrey, W.E. 1986. Birds of Canada. Nat. Mus.
Can. Bull. 203.
Godwin-Austen, H.H. 1874. Fourth list of birds
principally from the Naga Hills and Manipur,
including others from the Khasi Garo, and
Tipperah Hills./ Asiatic Soc. Bengal 43:151-
180.
Godwin-Austen, H.H. 1876. List of the birds
collected on the expedition into the Dafla
Hills, Assam, together with those obtained in
the adjacent Darrang Terai.y. Asiatic Soc.
Bengal 45:64-85.
Goeldi, E.A. 1894. Los Aves do Brasil. Livraria
Classica Alves, Rio de Janeiro.
Goeldi, E.A. 1897. Destruicao das garcas e
guaras. Bol. Mus. Paraense Hist. Nat. Ethnogr.
2:27-40.
Goeldi, E.A. 1902. Lista das Aves Amazonicas.
Bol. Mus. Paraense Hist. Nat. Ethnogr. 3:276-327.
Goeij, P. d., J. Kemper, and J. van Wetten. 1985.
De Lepelaar in het Waddengebied. Wadden
Bull. 20:85-37.
Goering, D.K. 1974. Organochlorine residues
and eggshell characteristics in three species of
ciconiiform birds. Diss. Abstr. Int. B Sci. Eng.
35:3111.
Gole, P. 1990. The status and ecological
requirements of Sarus Cranes: Phase II.
Unpublished report. Ministry of Environment
and Forests, Government of India, Delhi.
345
Gomez Dallmeier, F., and B.W. Cornaby. 1982.
Estudio de la comunidad de aves del
ecosistema de Punta Moron, Edo. Carabobo,
Venezuela. Pp. 821-348, in Adas del VIII
Congreso Latinoamericano de Zoologia, Merida.
Universidad de los Andes, Merida.
Gomez-Manzaneque, A. 1988. Tendidos electricos
en una zona de concentracion de Ciguena blanca
(Ciconia ciconia) en la provincia de Segovia.
Simposio sobre Ciguena Ibericas,
Guagelajaea, Abril 16-19, 1987.
Good, A.I. 1952. The birds of French Cameroon.
Mem. Inst. Franc. d'Afr. Noire 14:1-205.
Gooders, J. 1969. Birds of the World, vol. 1. IPC,
London.
Goodfellow, W. 1902. Results of an
ornithological journey through Colombia and
Equador.-part IV. Ibis (ser. 8) 2:59-67.
Goodman, S.M. 1988. A bird of antiquity.
Birder's World 2:26-29.
Goodman, S.M., and R.W. Storer. 1987. The
seabirds of the Egyptian Red Sea and adjacent
waters, with notes on selected Ciconiiformes.
Gerfaut 77:109-145.
Gordon, H.M. 1959. The Whale-headed Stork at
close range. Afr. Wildl. 13:83.
Gore, M.EJ. 1968. A check-list of the birds of
Sabah, Borneo. Ibis 110:165-196.
Gore, M.E.J., and Won P. O. 1971. The Birds of
Korea. Korean Branch of the Royal Asiatic
Society, Seoul, Korea. 450pp.
Goriup, P., and H. Schulz. 1990. Conservation
management of the White Stork: an international
opportunity. ICBP Study Report No. 37,
Cambridge, UK.
Gorski, W., R. Buskiewicz, E. Dabrowska, and
A. Grzeskowiak. 1977. Bocian bialy (Ciconia
ciconia) w dawnych powiatach chodzienskim i
slupeckim w Wielkopolsce w roku. [White
stork in Chodziez and Slupca Districts (Great
Poland) in 1974.] Notatki Orn. 18(3-4):95-99.
Gosper, D.G., S.V. Briggs, and S.M. Carpenter.
1983. Waterbird dynamics in the Richmond
Valley, New South Wales, 1974-1977. Aust.
Wildl. Res. 10:319-327.
Gotch, A.F. 1981. Birds- Their Latin Names
Explained. Blandford Press, Poole. 348pp.
Gould, J. 1838. A Synopsis of the Birds of Australia.
Pt. IV. J. Gould, London.
Gould, J. 1851. On a new and most remarkable
form in ornithology. Proc. ZooL Soc. Lond.
1851:1-2.
Gould, J. 1863. On Dr. Sclater's visit to
Amsterdam, Rotterdam, and Antwerp. Proc.
ZooL Soc. Lond. 1863:224-225,
Gould, J. 1865. Handbook to the Birds of Australia, 2
vols. Lansdowne Press, Melbourne.
Gradziel, T. 1968. Feeding of the Black Storks
Ciconia nigra together with the White Storks
Ciconia ciconia and herons Ardea cinerea. Notatki
Orn. 9:4.
Grafton, R.N. 1972. Survey of the Bald Ibis.
Fauna Flora (Pretoria) 23:16-19.
Grant, C.H.B. 1911. List of birds collected in
Argentina, Paraguay, Bolivia, and south
Brazil, with Field-notes. Ibis (ser. 9) 5:80-137,
317-349,459-478.
Grant, W.R.O. 1900. On the birds of Hainan.
Proc. ZooL Soc. Lond. 1900:457-504.
Gray, A.P. 1958. Bird hybrids: a checklist with
bibliography. Report No. 13 of the
Commonwealth Bureau of Animal Breeding
and Genetics, Edinburgh.
Gray, G.R. 1860. On additions to the menagerie.
Proc. ZooL Soc. Lond. 1860:184-185.
Gray, G.R. 1871. Hand-list of Genera and Species of
Birds. Part III. British Museum, London.
Greed, G. 1984. The White Stork (Ciconia
ciconia) —A review of the species with special
reference to breeding in the U.K., especially at
the Bristol Zoo. Pp. 4-10 in Partridge, J. (ed.),
Management of Cranes, Storks, and Ratites in
 Captivity, Proc. Symp. 9. Assoc. British Wild
Animal Keepers, Bristol, U.K.
Green, R.H. 1959a. The White Ibis in Tasmania.
Emu 59:58-60.
Green, R.H. 1959b. Straw-necked Ibis in
Tasmania. Emu 59:221.
Green, T. 1975. Roosting birds and the dangers
of a gas barge. Smithsonian 6:34-41.
Greene, E.R., W.W. Griffin, E.P. Odum, H.L.
Stoddard, and I.R. Tomkins. 1945. Birds of
Georgia, a preliminary check-list and Bibliography of
Georgia Ornithology. University of Georgia
Press, Athens, Georgia.
Greenewalt, G.H. 1962. Dimensional
relationships for flying animals. Smithson. Misc.
Collect. 144:1-46.
Green way, J.C., Jr. 1958. Extinct and Vanishing
Birds of the World. Spec. Publ. No. 13, Amer.
Comm. Intern. Wildlife Prot., New York.
518pp.
Greenway, J.C. 1967. Extinct and Vanishing Birds of
the World. Dover Publications, Inc., New York.
Greenwood, P.H. 1958. The Fishes of Uganda. The
Uganda Society, Kampala, 124pp.
Gregori, J. 1980. Black Stork in Slovenia.
Acrocephalus 1:25-26.
Greschik, J. 1922. Die Zunge von (Plegadis
falcinellus L.). Aquila 28:83-96.
Grieg-Smith, P.W. 1977. Breeding dates of birds
in Mole National Park, Ghana. Bull. Niger.
Orn. Soc. 1389-1393.
Griffith, R.E., V.L. Childs, and F.W. Cook.
1946. Roseate Spoonbill nesting on the Sabine
Refuge, Louisiana. Auk 63:259-260.
Grimmett, R. 1985. News and views: a protected
area network for Vietnam. Oriental Bird Club
Bull. 2:4.
Grimmett, R. 1985. News and views: more news
on Sumatran wetlands. Oriental Bird Club Bull.
2:5.
Grinnell, J., H.C. Bryant, and T.I. Storer. 1918.
The Game Birds of California. University of
California Press, Berkeley, CA.
Grischtenko, V.N. 1988. [Record of White Stork
(Ciconia ciconia) flock in summer.] Ornitologiya
23:207.
Grischtenko, V., and W. Boreiko. 1989.
Operation 'Storch' im Kiewer Gebiet der
UdSSR. Der Falke 36:99-102.
Grischtenko, V.N., V.Y. Boreiko, V.V.
Serebryakov, LA. Galinskaya. 1987. The
experience of conducting the popularization
campaign 'Leleka' in the Kiev Region. Pp.
101-102 (abstr.) in Ecological Education in the
U.S.S.R. UNESCO!ENEP International Congress
on Environmental Education and Training. Moscow
University Press, Moscow.
Griscom, L. 1932. The distribution of bird-life in
Guatemala. Bull. Am. Mus. Nat. Hist. 64:1-439.
Grote, H. 1932. Uber das Brutvorkommen von
Ciconia nigra (L.) in Suedafrika. Orn. Monatsber.
40:81-84.
Grote, H. 1954. Uber den
Schwarzschnabelstorch (Ciconia c. boyciana) im
unteren Amur-Gebiet. Vogelwarte 17:162-164.
Grummt, W. 1981. Breeding the Marabou Stork
Leptoptilos crumeniferus at Tierpark Berlin. Int.
Zoo Yb. 21:96-97.
Grunberg, W., and P.B. Diezel. 1965.
Metacercariencysten im Nervus vagus und
granulierende Encephalitis des Vaguskernes
bei einem Weiss-storch (Ciconia ciconia). Ada
Neuropathol. 4:336-339.
Grunberg, W., and E. Kutzer. 1964. Die
Pathologic verschiedener
Trematodeninfektionen bei Storchen (Ciconia
ciconia L., C. nigra L.). Z. Veterinaermed. Reihe B
11:712-727.
Gruson, E.S. 1976. Checklist of the Birds of the
World. Collins, London. 212pp.
Grussu, M. 1987. The breeding and wintering of
the Glossy Ibis, Plegadis falcinellus, and the
breeding of the Squacco Heron, Ardeola
ralloides, in Sardinia (Italy). Riv. Ital. Orn.
57:62-68.
Grzimek, B., E. Schuz, et al. 1968. Die Stoerche
und ihre Verwandten. Pp. 207—229 in Grzimeks
Tierleben, Band VII. Voegel I. Kindler Verlag,
Zuerich. 564pp.
Guichard, K.M. 1947. Birds of the innundation
zone of the River Niger, French Soudan. Ibis
89:450-489.
Guichard, K.M. 1950. A summary of the birds of
the Addis Abeba region, Ethiopia. J. E. Afr.
Nat. Hist. Soc. 19:154-181.
Guillet, A. 1978a. Distribution and conservation
of the Shoebill, (Balaeniceps rex) in the southern
Sudan. Biol. Conserv. 13:39-49.
Guillet, A. 1978b. Habitat and conservation of
the Shoebill Stork (Balaeniceps rex). Proc. Int.
Orn. Congr. 17:1367-1368.
Guillet, A. 1979. Aspects of the foraging
behaviour of the Shoebill. Ostrich 50:252-255.
Guillet, A. 1980a. Functional morphology and
feeding behaviour of Balaeniceps rex. Proc. Int.
Orn. Congr. 17:1367.
Guillet, A. 1980b. Habitat and conservation of
the Shoebill Stork (Balaeniceps rex). Proc. Int.
Orn. Congr. 17:1367-1368 (abstr.).
Guillet, A. 1984. Ecological and ethological
aspects of the nest of the Shoebill. Proc. Pan-
Afr. Orn. Congr. 5:231-236.
Guillet, A. 1987. Aspects of the evolution,
ecology and ethology of the Shoebill and their
bearing on a captive breeding program.
Pp.221-229 in J. Delacour (ed.), IFCB
Symposium: Breeding Birds in Captivity.
International Foundation for the Conservation
of Birds, Hollywood, California, USA.
Guillet, A., and T.M. Crowe. 1986. A
preliminary investigation of patterns of
distribution and species richness of southern
African waterbirds. S. Afr.J. Wildl. Res. 16:65-
81.
Guillet, A., W.S. Doyle, and H. Riither. 1985.
The combination of photogrammetry and
finite elements for a fine grained functional
analysis of anatomical structures.
Zoomorphology (Berl.) 105:51-59.
Gulliver, G. 1844. Additional measurements of
blood-corpuscles or red particles of mammalia
and birds. Proc. Zool. Soc. Lond. 1844:7-18.
Gulliver, G. 1875. Observations on the sizes and
shapes of the red corpuscles of the blood of
vertebrates, with drawings of them to a
uniform scale, and extended and revised tables
of measurements. Proc. Zool. Soc. Lond.
1875:474-495.
Gundlach, J. 1875. Neue Beitrage zur
Ornithologie Cubas.J. Orn. 23:293-407.
Gurney, J.H. 1899. On the comparative ages to
which birds live. Ibis (ser. 7) 5:19-42.
Gurtler, W.-D. 1989. Zu Besuch beim Kelaynak-
ein Nachruf auf die turkischen Waldrappen.
Zs. Kolner Zoo 32:149-156.
Gurung, K.K. 1983. Heart of theJungle. Andre
Deutsch, London.
Gyldenstolpe, N. 1945a. A Contribution to the
Ornithology of Northern Bolivia. Almqvist and
Amksells Boktryckeri AB, Stockholm.
Gyldenstolpe, N. 1945b. The bird fauna of Rio
Jurua in western Brazil. Kungl Svenska
Vetenskapsakademiens Handlingar 22.
Gyldenstolpe, N. 1955a. Birds collected by Dr.
Sten Bergman during his expedition to Dutch
New Guinea, 1948-1949. Arkiv Zoologi 8:183-
397.
Gyldenstolpe, N. 1955b. Notes on a collection of
birds made in the Western Highlands, Central
New Guinea, 1951. Arkiv Zoologi 8.
Gysels, H. 1968. Biochemical approach of the
central systematic position of the
ciconiiformes. Ardea 56:267—280.
Ha, B. 1991. Research in artificial rearing of
346
Oriental White Storks in captivity. Pp.207-212
in Coulter, M.C., Q. Wang, and C.S. Luthin
(eds), Biology and Conservation of the Oriental
White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Haagner, A., and R.H. Ivy. 1907. The birds of
the Albany Division, Cape Colony. J. S. Afr.
Orn. Union 3:76-116.
Haar, H. 1986. Projekt WeiBstorch: ein
Artenschutzprogramm zur Erhaltung des
WeiBstorchbestandes in der Steiermark.
Naturschutz Steiermark 129:19-22.
Haar, H. 1989a. Die Situation des WeiBstorchs
in der Steiermark. Vogelschutz Osterreich 4:20-
25.
Haar, H. 1989b. Zur Situation des Weiss-
Storches, Ciconia ciconia L., in Osterreich. Pp.
65-68 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weijtstorch: Status and Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985. Schriftenr. DDA, Braunschweig,
Germany.
Haar, H. 1990. WeiBstorch (Ciconia ciconia) aus
den Vojvodina (Jugoslawien) bruetet in
Apelton, Burgenland. Vogelwarte 35:243.
Haarhoff, PJ. 1988. A new fossil stork (Aves,
Ciconiidae) from the late Tertiary of
Langebaanweg, South Africa. Ann. S. Afr. Mus.
97:297-313.
Haas, G. 1955. Ueber Storchkaempfe, ihre
Deutung und Bedeutung nach Beispielen 1955
am Federsee. Aus der Heimat
63(11/12):208-218.
Haas, G. 1963. Nestwechsel, Gelege-
Uebernahme, Zusatz- und Nachgelege bei
weiblichen WeiBstoerchen. Vogelwarte 22:100-
109.
Haas, G. 1964. Horst- und Partnerwechsel eines
mannlichen WeiBstorchs innerhalb einer
Brutzeit.J^. Ver. Vaterl. Naturkde. Wurttemberg
118/119:382-385.
Haas, G. 1966. Jungenverluste bei
WeiBstorch-Gehecken mit zweierlei
Altersgruppen. Vogelwarte 23:300-305.
Haas, G., and W. Haas. 1965. Schwere
Erkrankung eines Weiss-storchs durch
Behinderung der Gewollabgabe. J/L Ver. Vaterl.
Naturk. Wurtt. 120:292-295.
Hachisuka, M. 1931-1932. The Birds of the
Philippine Islands, vol. 1. H.F. and G. Witherby,
London.
Hachisuka, M., and T. Udagawa. 1951.
Contribution to the ornithology of Formosa,
Part II. Q, J. Taiwan Mus. (Taipei) 4:1-180.
Haedo-Rossi, J.A. 1969. Notas ornitologicas v
observaciones sobre la Ciguena Euxenura
maguari (Gmelin). Acta Zool. Lilloana 25:19-42.
Haffenden, A. 1981. Arboreal foraging by a
Straw-necked Ibis (Threskiornis spinicollis).
Sunbird Il(38c4):76.
Hagmann, G. 1906. Ornithologisches von der
Insel Mexiana, Amazonenstrom. Ornithol.
Monatsber. 14:107-111.
Hagmann, G. 1907. Die Vogelwelt der Insel
Mexiana, Amazonenstrom. Zool. Jahrb. Abt.
Syst. 26:11-62.
Hailman, J.P. 1959. Consolidation of northward
extension of the Glossy Ibis's breeding range.
Bird-Banding 30:231-232.
Hails, C., and O. Mueller. 1985. Surveying and
ringing in Singapore. Interwader Ann. Rep.
1984:119-126.
Hall, B.P. 1974. Birds of the Harold Hall Australian
Expeditions 1962-1970. Trustees Brit. Mus.
(N.H.), London. 396pp.
Hall, J.V., and W.C. Cone. 1970. Wood Ibis
found nesting in Okeefenokee Refuge. Oriole
35:14.
Hall, M.R., E. Gwinner, and M. Bloesch. 1987.
Annual cycles in moult, body mass, luteinizing
hormone, prolactin and gonadal steroids
 during the development of sexual maturity in
the White Stork (Ciconia ciconia). J. Zool. Lond.
211:467-486.
Hall, R.G. 1964. Roseate Spoonbills in numbers
near Sebastian Inlet. Fla. Nat. 37:121.
Hallinan, T. 1924. Notes on some of the Panama
Canal Zone birds with special reference to
their food. Auk 41:304-326.
Hamel, H.D. 1975. Ein Beitrag zuer
Populationsdynamik des Waldrapps Geronticus
eremita(L., 1758). Vogelwelt 96:2 13-221.
Hamel, P.B. 1977. The Wood Stork in South
Carolina, a review. Chat 41:24-27.
Hamernick, J. 1989. An ibis in Marshall County.
Loon 61:201.
Hamerton, A.E. 1943. Report on the deaths
occurring in the Society's Gardens during the
year 1941. Proc. Zool. Soc. Lond. 112:120-136.
Hamilton, E. 1874. [Untitled.] Proc. Zool. Soc.
Lond. 1874:306-307.
Hamling, H.H. 1953. Observations on the
behaviour of birds in Southern Rhodesia.
Ostrich 24:9-16.
Hammatt, R.B. 1981. Reproductive biology in a
Louisiana heronry. M.S. Thesis, Louisiana
State University, Baton Rouge, Louisiana.
Hammerschimdt, R. 1968. Zum einflug von
Braunsichlern (Plegadis falcinellus) im Hebst
1967 nach Westfalen und Westneidersachen.
Orn. Mitt. 20:60.
Hanagarth, W., and M. Octavio Riberia. 1985.
Los Ciconiidae de Bolivia. Ecol. Bolivia 6:73-
81.
Hancock, J. 1969. America's stork. Animals
(Lond.) 12:308-309.
Hancock, J. 1984. The Birds of the Wetlands.
Groom Helm, London.
Hancock, J. 1989. Extinction stalks the storks of
Asia. World Birdwatch 11(1):1.
Hancock, J. 1990. Status and conservation needs
of larger water birds of South Asia [abstract].
P. 46 in Seminar on Wetland Ecology and
Management. Bombay Nat. Hist. Soc.,
Bharatpur, India.
Hancock, J., andj. Kushlan. 1984. The Herons
Handbook. Groom Helm, London, and Harper
& Row, New York.
Hanebrink, E.L., and R. Cochran. 1966. An
inland nesting record of the Glossy Ibis,
Plegadis falcinellus, in Arkansas. Auk 83:474.
Hanmer, D.B., 1989. Comments on the Shoebill
in Malawi. Nyala 14:46.
Hanmer, D.B., and M. Roseveare. 1989. First
record of the Shoebill Balaeniceps rex in Malawi.
Scopus 12:92-93.
Haraszthy. L. 1989. Die Situation des Weiss-
Storchs in Ungarn. Vogelschutz Osterreich 4:18-
19.
Harber, D.D. 1955. Special review of the Birds of
the Soviet Union, Vol. 2. British Birds 48:268-
276.
Harington, H.H. 1909-1910. A list of the birds of
the Bhamo District, Upper Burma. J. Bombay
Nat. Hist. Soc. 19:299-313.
Harley, D. 1990. Annual performance report:
Wood Stork surveys. Unpublished report of
Georgia DNR Project No. E-l-14, Study XL
Georgia Department of Natural Resources.
Harris, J. 1988. The storks of Wat Phai Lorn.
i.c.f. Bugle 14:1
Harrison, E.G. 1933. The Bald Ibis (Comatibis
eremita (L.)). Oologists' Rec. 13:74.
Harrison, C.J.O. 1974. A re-examination of
material of the extinct Marabou Stork,
Leptoptilos falconeri: with descriptions of some
new species. Bull. Br. Orn. Club 94:42-49.
Harrison, C J.O. 1975. A re-examination of the
extinct Pleistocene stork Palaeociconia australis.
Bull. Br. Orn. Club 95:94-96.
Harrison, C.J.O. 1980. Fossil birds from
Afrotropical Africa in the collection of the
British Museum (Natural History) . Ostrich
51:92-98.
Harrison, C. 1982. An Atlas of the Birds of the
Western Palearctic. Princeton University Press,
Princeton, New Jersey.
Harrison, C.J.O., and C.A. Walker. 1971. A new
ibis from the lower Eocene of Britain. Ibis
113:367-368.
Harrison, H.H. 1975. A Field Guide to Birds' Nests.
Houghton Mifflin Company, Boston.
Harrison, J.W. 1978. A Field Guide to the Nests,
Eggs, and Nestlings of North American Birds.
Collins, New York.
Hartert, E. 1902. Some further notes on the birds
of northwest Ecuador. Novit. Zool. 9:599-617.
Hartert, E. 1912-1921. Die Voegel der
Palaearktischen Fauna, vol. 2. R. Friedlaender,
Berlin.
Hartert, E. 1913. Expedition to the central
western Sahara. IV. Birds. Novit. Zool. 20:37-
76.
Hartert, E., and F.C.R. Jourdain. 1923. The
hitherto unknown birds of Morocco. Novit.
Zool. 30:91-146.
Hartert, E., and W.R. Ogilvie-Grant. 1905. On
the birds of the Azores. Novit. Zool. 12:80-128.
Hartert, E., and S. Venturi. 1909. Notes sur les
oiseaux de la Republic Argentine. Novit. Zool.
16:159-267.
Hartley, A., P. Tongue, E.M. van Zinderen
Bakker, J.M. Winterbottom, and M.G.
Winterbottom. 1968. Breeding of Glossy Ibis
on Berg River, Cape Province. Ostrich 39:39-
40.
Hartman, F.A. 1955. Heart weight in birds.
Condor 57:22 1-238.
Hartman, F.A. 1961. Locomotor mechanisms of
birds. Smithson. Misc. Collect. 143:1-91.
Hartmann, F. A. 1968. Africa's most beautiful
stork: the Saddle-bill. Africana 3(8): 16-1 7.
Hartman, F.A., and R.H. Albertin. 1951. A
preliminary study of the avian adrenal. Auk
68:202-209.
Harvey, W.G. 1972. Woolly-necked Stork Ciconia
episcopus near Dar es Salaam. East Afr. Nat.
Hist. Soc. Bull. Oct:\69.
Harvey, W.G., and K.M. Howell. 1987. Birds of
the Dar Es Salaam area, Tanzania. Gerfaut
77:205-258.
Harwin, R.M. 1970. Some aspects of the
recovery of ringed White Storks in Rhodesia.
Honeyguide 62: 1 5—16.
Harwin, R.M. 1971. White Stork: longevity
record. Ostrich 42:81.
Haucke, H.H., and W.H. Kiel, Jr. 1973. Jabiru
in south Texas. Auk 90:675-676.
Haverschmidt, F. 1935. Beobachtungen in der
Loflerkolonie im Zwanenwater. Beitr.
Fortpflanzungsbiol. Vogel. 11:1-3.
Haverschmidt, F. 1938. Spoonbills in Holland.
Bird Lore 40:325-327.
Haverschmidt, F. 1949. The Life of the White Stork.
E.J. Brill. Leiden, Holland.
Haverschmidt, F. 1X950. Bemerkungen uber den
Weissen Storch (Ciconia ciconia (L.)) im
nordlichen Zugscheidegebiet. Orn. Beob.
47:73-79.
Haverschmidt, F. 1952. The White Stork, Ciconia
ciconia, in South Africa. Ostrich 23:125.
Haverschmidt, F. 1955a. The White Stork.
Audubon Mag. 57:32-34.
Haverschmidt, F. 1955b. List of the birds of
Surinam. Publ. Found. Sci. Res. in Surinam
and Netherlands Antilles, Utrecht No. 13.
153pp.
Haverschmidt, F. 1956. Das Wenden der Eier
beim WeiBstorch. Vogelwelt 77:1 18.
Haverschmidt, F. 1962. Notes on some Suriname
breeding birds (II). Ardea 50:173-179.
Haverschmidt, F. 1966. Naschrift/Laatste
bericht.Ardea 54:12_13.
Haverschmidt, F. 1967. De broedkolonies van de
347
Rode Ibis (Eudocimus ruber) in Suriname in
1966. Ardea 55:141-143.
Haverschmidt, F. 1968. Birds of Surinam. Oliver
and Boyd, Edinburgh, London.
Haverschmidt, F. 1969. The Roseate Spoonbill
breeding in Surinam. Auk 86:131.
Havre, G.C.M. v. 1928. Les Oiseaux de la Fame
Beige. Lamertin, Bruxelles.
Haweis, S. 1929. The Black Stork. Nature Mag.
13:35-37.
Hawkins, A. 1986. Northward migration 1986-
south and eastern peninsular Malaysia.
Interwader Newsletter 7:5.
Hawkins, A., and M. Silvius. 1986. West Coast
Peninsular Malaysia. Interwader Newsletter 7.
Hays, H. 1967. The adaptive ibis. Nat. Hist.
66:32-33.
He, B. 1981. [Training-conditioned reflex and
behaviour of people recognition by White
Storks.] Chinese J. Zool. 2:11.
He, B. 1987. [Preliminary research on hand-
rearing Oriental Ibis.] Chinese Wildl. 2:25.
He, B. 1988. [Preliminary research on hand-rearing
White Storks.} Symposium of the Ornithological
Society of China IV.
Healy, M. 1991. Maguari Storks hatch at
Discovery Island, Florida. Spec. Group Storks,
Ibises, Spoonbills Newsletter 4(l):5-6.
Hebard, F.V. 1941. Winter birds of the
Okeefenokee and Coleraine. Georgia Soc. Nat.
Bull. 3:13.
Hebard, F.V. 1950. Glossy Ibis breeding in
Georgia. Oriole 15:9-10.
Hecht, T. 1981. A bird paradise in jeopardy.
Bokmakierie33:\2-\5.
Heckenroth, H. 1985. Atlas der Brutvoegel
Niedersachsens 1980. Hannover: Naturschutz
und Landschaftspflege in Niedersachsen Heft
14.
Heckenroth, H. 1989. Zur Situation des Weiss-
Storchs in der Bundesrepublik Deutschland.
Pp. 60-61 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), WeiBstorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Heckenroth, H. 1990. WeiBstorch, Ciconia ciconia,
Brutbestand 1971, 1988 and 1989.
Neldersachsen und Bremer, Nordrhein-
Westfalen, Hamburg suedlich Elbe/
Suederclbe. Niedersaechs. Landesver
Waltungsamt-Fachbehoerde Jur Naturschutz,
Hannover.
Heckenroth, H., and E. Schuz. 1970. Funde in
Europa beringter WeiBstorche im Orient
ostlich der Schmalfront. Zool. Abh. (Dres.)
31:193-203.
Heckenroth, H., M. Frantzen, R. Berndt, H.
Ringleben, and A. Festetics. 1976. Rote Liste
der in Niedersachsen gefahrdeten. 2. Fassung,
Stand 1.1.1976.
Hediger, H. 1947. 1st das tierliche Bewusstein
unerforschbar? Behaviour 1:130-137.
Hediger, H. 1951. Observations sur la Psychologie
Animale dans'les Pares Nationaux du Congo Beige.
Institut des Pares Nationaux du Congo Beige,
Brussells.
Hediger, H. 1955. Studies of the Psychology and
Behaviour of Captive Animals in Zoos and Circuses.
Criterion, New York.
Heeg, J., and C.M. Breen. 1982. Man and the
Pongolo floodplain. S. Afr. Nat. Sci. Progr. Rep.
56:1-117.
Heer, E., and I. Schoch. 1952. Vom Weissen
Storch in den deutschen Kolonien Sud-
Bessarabiens. JA. Ver. Vaterl. Naturk. 107:72-
88.
Heiden, J.T. v.d. 1973. Openbill Storks nesting
near Salisbury. Honeyguide 76:22-25.
Heiden, J.T. v.d. 1974. Openbill Storks nesting
near Salisbury: a postscript. Honeyguide 77:33-
36.
Heim de Balsac, H. 1924. L'Ibis chauve,
Comatibis eremita (L.), en Algeria. Rev. Fr, Orn.
8:469-474.
Heim de Balsac, H. 1925. Quelques oiseaux
particulierment interessants de 1'Algerie et du
Sahara. Rev. Fr. Orn. 9:189-193.
Heim de Balsac, H. 1952a. Rythme sexuel et
fecondite chez les oiseaux du nord-ouest de
1'Afrique. Alauda 20:213-242.
Heim de Balsac, H. 1952b. Considerations sur
une biocenose constituee autour d'un nid de
cigogne, Ciconia ciconia, en Lorraine. Alauda
20:144-156 (with English Summary).
Heim de Balsac, H. 1964. Les recentes
acquisitions faunisitiques effectuees par R. de
Naurois sur la cote Atlantique, du Maroc au
golfe de Guinee. Alauda 32:245-249.
Heim de Balsac, H., and T. Heim de Balsac.
1951. Les migrations des oiseaux dans 1'ouest
du continent africain. Alauda 19:157-171.
Heim de Balsac, H., and N. Mayaud. 1962. Les
Oiseaux du Nord-Ouest de 1'Afrique. Editions Paul
Lechevalier, Paris. 486pp.
Heimendahl, A. 1985. Untersuchungen zur
Nestlingsentwicklung des Scharlachsichlers
(Eudocimus ruber). Thesis, University of
Hamburg.
Heinrichs, J. 1986. Spoon feeder. Natl Wildl. 24
(Jun/Jul):38-41.
Heinroth, K. 1963. Beobachtungen zur
Fluchtdistanz. Z. Tierpsychol. 20:624-626.
Heinroth, O. 1905. Bericht uber die 54.
Jahresversammlung. J. Fr. Ornith., 53:249-
261.
Heinroth, O. 1922. Die Beziehungen zwischen
Vogelgewicht, Eigewicht, Gelegegewicht und
Brutdauer.J. Orn. 70:172-285.
Heinroth, O. 1924. Lautausserungen der Voegel.
J. Orn. 72:223-244.
Heinroth, O., and M. Heinroth. 1924-1931. Die
Voegel Mitteleuropas. 4 Vols. Bermuhlerverlag,
Berlin-Lichterfelde.
Heinze,J., and N. Krott. 1979. Zur Vogelwelt
Marokkos (2). [The bird world of Morocco 2.]
Vogelwelt 100:225-227.
Heinzel, H., and D. Martinoles. 1988. Nouvelle
nidification de 1'Ibis falcinelle Plegadis
falcinellus en France. [Another case of the
Glossy Ibis Plegadis falcinellus nesting in
France.] Alauda 56:429-430.
Heinzman, G., and D. Heinzman. 1965. Nesting
of the Wood Stork. Nat. Hist. 74:30-35.
Held, M.M. 1990. Status and conservation of the
Scarlet Ibis in Suriname. Pp. 100-106 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Hellebrekers, W.PJ. 1942. Revision of the
Penard oological collection from Surinam.
Zoo/. Meded. 24:240-275.
Hellebrekers, W.PJ. 1967. Gegevens over de
eieren van min of meer zeldzame of schaarse
inlandse broedvogels in Nederlandse collecties.
Hellebrekers, W.PJ., and A. Hoogerwerf. 1967.
A further contribution to our oological
knowledge of the island of Java (Indonesia).
Zoo/. Verh. (Leiden) 88: 1-164.
Heller, E. 1932. A record of the first nesting of
the American Wood Stork in captivity. Bull.
Wash. Park Zoo/. Soc. Milwaukee 3:2-4.
Hellmayr, C.E., and B. Conover. 1948. Birds of
the Americas. Field Mus. Nat. Hist. Publ. Zoo/.
Ser. 13:1-434.
Helps, M J. 1968. Storks over Istanbul. Animals
(Lond.) 10:468-471.
Hemetsberger, J. 1989. Bestandsentwicklung und
Derzeitige verbreitung de Schwarzstorches
(Ciconia nigra) in Oberoesterreich. [Population
dynamics and present distribution of the Black
Stork (Ciconia nigra) in Upper Austria.] Stapfia
20:119-128.
Hemke, E. 1988. Tagung zum Schutz des
WeiBstorches. Falke 6:199.
Hemke, E. 1969. Concentration of storks in the
case of a widespread conflagraton. Falke
Monatsschr. Orn. Vivarienkunde Ausg A 16:13.
Hemming, F. 1952. Proposed correction...for the
generic name 'Balaeniceps'. Bull. Zoo/. Nomencl.
9:85-86.
Hemmingsen, A.M. 195la. Observations on the
migration of the Eastern White Stork (Ciconia
ciconia boyciana Swinhoe). Proc. Int. Orn. Congr.
10:351-353.
Hemmingsen, A.M. 1951b. Observations on
birds in North Eastern China, especially the
migration at Pei-Tai-Ho Beach. Spolia Zoo/.
Mus.Haun. 11:1-227.
Hemmingsen, A.M., and J.A. Guildal. 1968.
Observations on Birds in North Eastern
China. II. Spolia Zoo/. Mus. Haun. 28:1-326.
Henderson, E.G. 1981. Behavioral ecology of the
searching behavior of the White Ibis
(Eudocimus albus). M.S. Thesis, University of
South Carolina, Columbia, South Carolina.
Hendren, L., and B.R. Chamberlain. 1955.
Remarkable flight of the Wood Ibises. Chat
19:85.
Henny, C J., and G.B. Herron. 1989. DDE,
selenium, mercury, and White-faced Ibis
reproduction at Carson Lake, Nevada. J.
Wildl. Manag. 53:1032-1045.
Henny, C J., LJ. Blus, and C.S. Hulse. 1985.
Trends and effects of organochlorine residues
on Oregon and Nevada wading birds 1979-83.
Colonial Waterbirds 8:117-128.
Henry, G.M. 1955. A Guide to the Birds of Ceylon.
Oxford Univ. Press, London.
Henry, G.M. 1971. A Guide to the Birds of Ceylon,
2nd edn. K. V. De Silva and Sons, Kandy, Sri
Lanka.
Herbert, E.G. 1926. Nests and eggs of birds of
central Siam. J. Siam Soc., Nat. Hist. Suppl.
6:323-356.
Herklots, G.A.C. 1961. The Birds of Trinidad and
Tobago. Collins, London.
Hernandez, R. 1953. Scarlet Ibis come home
again. Trinidad Guardian Aug. 7.
Heron, S J. 1973. Birds of the Orang District,
NSW. £>m/73:108.
Heron, S J. 1970. The birds of middle Swan,
Western Australia. Emu 70:155-158.
Herroelen, P. 1955. Notes sur quelques nids et
oefs inconnus d'oiseaux African observes au
Congo Beige. Rev. Zoo/. Bot. Afr. 52:185-192.
Herzig, A. 1989. Die WeiBstorch-
Aufzuchtstation Schwarzach 1989. Gejiederte
Welt 113:342-344.
Heuglin, M.T. von. 1869-1873. Ornithologie
Nordost-Afrika's der Niquellen- Kuesten-
Gebiete des Rothen und der nordlichen
Somali-Landes. T. Fischer, Cassell, Germany.
Heurn, W.C. v. 1912. Enkele maanden
ornithologiseeren in Suriname, 29 Mei-22
October 191 \.Jaarb. Club van Nederl.
Vogelkundigen 2:69-108.
Hewetson, C.E. 1956. Observations on the bird
life of Madhya Pradesh. J. Bombay Nat. Hist.
5-0(7.37:81-95.
Hewlett, J. 1960. Bald Ibises sheltering in a cave
from inclement weather. Ostrich 31:29.
Hey, T. 1974. The storks that lost their way.
Honeyguide 77:36-37,41.
Hiatt, B.C. 1933. Avocets and spoonbills on
Merritts Island, Florida. Auk 50:100.
Higgins, J.C. 1934. The game birds and animals
of the Manipur State with notes on their
numbers, migration and habits, part IV. J.
Bombay Nat. Hist. Soc. 37:81-95.
Higuchi, Y. 1976. A Japanese White Stork
348
(Ciconia ciconia boyciana) obtained in Mie
Prefecture. Misc. Rep. Yamashina Inst. Orn. 8:79-
81.
Hilderbrandt, H. 1927. Wie entstand Bechsteins
Waldrappbild./ Orn. 75:519-522.
Hill, A., and A. Walker. 1979. A fossil Marabou
(Aves: Ciconiidae) from the Miocene Ngorora
Formation, Baringo District, Kenya. Neth. J.
Zoo/. 29:215-220.
Hill, W.C.O. 1943. Intergeneric hybrid storks
hatched at semi-liberty. Avic. Mag. 8:141-144.
Hill, W.C.O., and C J. Skead. 1952. On terminal
claws on the manual digits in Ardeiform birds.
Ibis 94:62-67.
Hilty, S.L., and W.L. Brown. 1986. A Guide to the
Birds of Colombia. Princeton University Press,
Princeton, New Jersey.
Himmatsinhji, M.K. 1985. The Black Stork in
Kutch: old record confirmed./ Bombay Nat.
Hist. Soc. 82:403.
Himmatsinhji, M.K. 1989. Occurrence of Ciconia
ciconia Gruidae and breeding of
Phoenicopteridae in Kutch, Gujarat. J. Bombay
Nat. Hist. Soc. 86:443-444.
Hintz, W. 1856. Auszuge aus meinem
ornithologischem Tagebuche. Naumannia 6:18—
27.
Hirsch, U. 1973. Project 945. Conservation of the
Bald Ibis. World Wildl. Yb. 1972-1973:151-
152.
Hirsch, U. 1974. Project 945. Bald Ibis
conservation. WWF Yb. 1973-1974:198.
Hirsch, U. 1975. Project 945. Bald Ibis
conservation. WWF Yb. 1974-1975:196.
Hirsch, U. 1976a. Project 1288. Bald Ibis,
Morocco-Survey. WWF Yb. 1975-1976:77-78.
Hirsch, U. 1976b. Beobachtungen am Waldrapp
Geronticus eremita in Marokko und Versuch zur
Bestimmung der Alterszusammensetzung von
Brutkolonien. Orn. Beob. 73:225-235.
Hirsch, U. 1977a. Aufruf zur Zusammenarbeit
'Zum Scutz des Waldrapps Geronticus eremita'.
Vogelwelt 98:40.
Hirsch, U. 1977b. Co-operation invited on the
protection of the Bald Ibis Geronticus eremita.
Bull.Br. Orn. Club 97:72.
Hirsch, U. 1978a. Artificial nest ledges for Bald
Ibises. Pp. 61-69 in Temple, S.A. (ed.),
Endangered Birds: Management Techniques for
Preserving Threatened Species. University of
Wisconsin Press, Madison, Wisconsin.
Hirsch, U. 1978b. Zum Schutz des Waldrapps
(Geronticuseremita). J. Orn. 119:465-466.
Hirsch, U. 1979a. Hilfe fur den Waldrapp. Wir
Vogel 11:12-15.
Hirsch, U. 1979b. Studies of west Paleartic birds
183, Bald Ibis. British Birds 72:313-325.
Hirsch, U. 1979c. Protection of Geronticus eremita.
Bull. Br. Orn. Club 99:39.
Hirsch, U. 1980a. Der Waldrapp Geronticus
eremita, ein Beitrag zur Situation in seinem
ostlichen Verbreitungsgebiet. [The Waldrapp
Ibis Geronticus eremita: the situation in the
eastern part of its distribution area.] Vogelwelt
101:219-236.
Hirsch, U. 1980b. Project 1631. Waldrapp or
Bald Ibis. WWF Yb. 1979-1980:130.
Hirsch, U. 198la. Project 1631. Waldrapp Ibis.
WWF 70.1980-1981:213.
Hirsch, U. 1981b. Morocco plans reserve for
Waldrapp Ibis. WWF Monthly Rep.
January :21-23.
Hirsch, U. 1981c. Captive breeding for
Waldrapp Ibis. WWF Monthly Rep.
January: 17-19.
Hirsch, U. 198Id. Morocco declares reserve for
Waldrapp Ibis. WWF Monthly Rep. July: 191-
193.
Hirsch, U. 1981e. Conservation Project Bald
Ibis: Waldrapp, Geronticus eremita, 1981.
Unpublished report, WWF/IUCN, Morges
Switzerland.
Hirsch, U. 1982a. Waldrapp Ibis, Geronticus
eremita. Unpublished report, WWF
International, Gland, Switzerland.
Hirsch, U. 1982b. Morocco Project 1631.
Waldrapp Ibis. WWF Yb. 1982:211-212.
Hirsch, U., and A. Schenker. 1977. Der
Waldrapp (Geronticus eremita).
Freilandbeobachtungen und Hinweise fur eine
artgemasse Haltung. Zeitschr. Koelner Zoo
20:3-11.
Hislop, G., and C.James. 1990. Status and
conservation of the Scarlet Ibis in Trinidad
and Tobago. Pp. 115-123 in Frederick, P.C.,
L.G. Morales, A.L. Spaans, and C.S. Luthin
(eds), The Scarlet Ibis (Eudocimus ruber):
Status, Conservation, and Recent Research, Proc. 1st
International Scarlet Ibis Conservation
Workshop, Caracas, Venezuela. IWRB
Special Publication 11, Slimbridge, Glos, UK.
Hitchins, P.M. 1974. Breeding localities of the
Woollynecked Stork in Zululand game
reserves. Lammergeyer 21:47-49.
Hladik, B. 1989. Bestandsaenderungen des
WeiBstorchs im Nordosten des
bohmischmahrischen Huegellandes. Pp. 77-80
in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weiflstorch: Status undSchutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Hobbs, J.N. 1958. Herons and ibis sun-bathing.
Emu 58:286-287.
Hocke, A. 1904. Warum wirft der Storch seine
Jungen hinaus? Sanct Hubertus 22:280.
Hodgson, M.E.,J.R.Jensen, H.E. Mackey,Jr.,
and M.C. Coulter. 1987. Remote sensing of
wetland habitat: a Wood Stork example.
Photogramm. Eng. Rem. Sensing 53:1075-1080.
Hodgson, M.E.,J.R.Jensen, H.E. Mackey,Jr.,
and M.C. Coulter. 1988. Monitoring Wood
Stork foraging habitat using remote sensing
and geographic information systems.
Photogramm. Eng. Rem. Sensing 54:\6Q\-\6Q7.
Hoefler, J.E. 1979. White Ibis. Passenger Pigeon
41:43.
HoevellJJ.L.D. (Baron) van. 1911.
Aanteekeningen over de reticulaire cellen bij
eenige verschillende vertebraten. Versl. Gewone
Vergad. Akad. Amst. 19:988-1006.
Hofer, H. 1954. Neuere untersuchungen zur
Kopfmorphologie der Voegel. Proc. Int. Orn.
Congr. 11:104-115.
Hoffman, G. 1963. Drei Storche auf einem Horst.
0m.Affcr.43:120.
Hoffman, R., R. Faust, G. Hoffman-Fezer, and
U. Weinand. 1974. Karyotypen von Kagu,
Klunkerkranich und Schuhschnabel. Zoo/.
Garten (N.F.) 44:349-356.
Hoffman, R.H. 1971. Sighting of a Glossy Ibis in
Jackson County. Jack-pine Warbler 4:9.
Hoffman, T.W. 1984. National Red Data List of
Endangered and Rare Birds of Sri Lanka. Ceylon
Bird Club and Wild Life and Nature
Protection Society of Sri Lanka, Colombo.
Hoffman, T.W. 1985. The 2nd duck count in Sri
Lanka (Mid-January 1984). Loris 17:19-24.
Hoffmann, E. 1978. Ten thousand Scarlet Ibis.
Americas 30:45-50.
Hofmeyr, J. 1990. White storks overwintering.
/Vowro/u 195:11-12.
Hogg, P., PJ. Dare, and J.V. Rintoul. 1984.
Palaearctic migrants in the central Sudan. Ibis
126:307-331.
Hoier, R. 1955. A Travers Plaines et Volcans au Pare
National Albert, 2nd edn. Inst. des Pares
Nationaux du Congo Beige, Bruxelles.
Holdaway, R.N. 1980. Royal Spoonbills nesting
near Blenheim. Notornis 27(2): 168-169.
Holdsworth, E.W.H. 1872. Catalogue of the
birds found in Ceylon; with some remarks on
their habits and local distributions, and
descriptions of two new species peculiar to the
island. Proc. Zoo/. Soc. Lond. 1872:404-483.
Holland, A.H. 1892. Short notes on the birds of
the Estancia Espartilla Argentine Republic.
7£«(ser. 6)4:193-214.
Hollom, P.A.D. 1959. Notes from Jordan,
Lebanon, Syria and Antioch. Ibis 101:183-
200.
Holmes, D.A. 1969. Bird notes from Brunei:
December 1967-September 1968. Sarawak Mus.
J. 17:399-402.
Holmes, D.A. 1977. Faunistic notes and further
additions to the Sumatran avifauna. Bull. Br.
Om. Club 97:68-71.
Holmes, D.A., and K. Burton. 1987. Recent
notes on the avifauna of Kalimantan. Kukila
3:2-7.
Holstein, V. 1929. Skehejren, Platalea leucerodia,
[sic] som ynglende i Danmark. Dan. Orn.
Foren. Tidsskr. 22:111-118.
Holt, E.G. 1929. In the haunts of the Wood Ibis.
Wilson Bull. 41:2-18.
Holt, E.G. 1933. Definite breeding of the Glossy
Ibis in Louisiana. Auk 50:351-352.
Holt, E.G., and G.M. Sutton. 1926. Notes on the
birds observed in southern Florida. Ann.
Carnegie Mus. 16:409-439.
Holyoak, D. 1970. Comments on the
classification of Old World Ibises. Bull. Br.
Orn. Club 90:67-73.
Homonnay, F. 1964. Quantitative
Storchaufnahme in Ungarn und in den
anstossenden Gebieten im Jahre 1941. Aquila
69/70:83-102.
Hommonay, N. 1959. The spoonbill colonies on
the fishponds of the Hortobagy. Aquila 66:307.
Hoogerwerf, A. 1935. Broedende Whitte Ibissen
in West-Java. Tropische Nat. 24:163-171.
Hoogerwerf, A. 1936a. On the nidification of
some Javan birds. Bull. Raffles Mus. 12:118-
124.
Hoogerwerf, A. 1936b. Voorkomen van Nycticorax
caledonicus (Gm.), Plegadis falcinellus subspec. en
Phalacrocorax ater territori (Mathews) als
broedvogel in Oost-Java. Ardea 25:170-171.
Hoogerwerf, A. 1936c. Met de camera naar de
Nimmerzatten. Tropische Nat. 25:83-96.
Hoogerwerf, A. 1937. Uit het leven der Witte
Ibissen Threskiornis aethiopicus melanocephalus.
Limosa W(4):\37-\46.
Hoogerwerf, A. 1948. Contribution to the
knowledge of the distribution of birds on the
island of Java: Treubia 19:83-137.
Hoogerwerf, A. 1949. Een Bijdrage tot de Oologie
van het Eilandjava. Koninklijke Plantentuin van
Indonesee, Buitenzorg, Java.
Hoogerwerf, A. 1951. Nieuwe bijzonderheden
over het voorkomen van de lepelaar, Platalea
leucorodia regia Gould, in west-Java. Limosa
24:91-99.
Hoogerwerf, A. 1952. De lepelaars (Platalea
leucorodia regia Gould) van Pulau Dua (West-
Java) gedurende het broedseizoen 1952. Limosa
25:118-131.
Hoogerwerf, A. 1953. Zwarte ibissen, Plegadis
falcinellus .peregrinus (Bp.), in het vogelreservaat
Pulau Dua in dejaren 1951 en 1952. Limosa
26:20-30.
Hoogerwerf, A. 1964. On birds new for New
Guinea or with a larger range than previously
known. Bull. Br. Orn. Club. 84:70-77.
Hoogerwerf, A., and G.F.H.W. Rengers Hora
Siccama. 1937. De avifauna van Batavia en
omstreken. Ardea 26:1 16-159.
Hoogstraal, H., M.N. Kaiser, and H.E.
McClure. 1975. The subgenus Persicargas
(Ixodoidea: Argasidae: Argas) 20. A. (P.)
robertsi parasitizing nesting wading birds and
domestic chickens in the Australian and
Oriental Regions, viral infections, and host
migration./ Med. Entomol. 11:513-524.
Hopkins, M.N. Jr. 1971. Another breeding
record for the White Ibis in mid-south
Georgia. Oriole 36:8-9.
349
Hopkins, M.N. and R.L. Humphries. 1983.
Observations on a Georgia Wood Stork
nesting colony. Oriole 48:36-39.
Hopkinson, E. 1926. Records of Birds Bred in
Captivity. Witherby, London.
Hoppe, D. 1990. Nun ist den Waldrapp aus
Eurasien endgueltig verschwunden. Gef. Welt
114:36.
HopwoodJ.C. 1909-1910. Occurrence of the
spoonbill (Platalea leucorodia) in lower Burma.
J. Bombay Nat. Hist. Soc. 19:261.
Hopwood, J.C. 1921. Notes on some nests
recently found in South-Tenasserim./ Bombay
Nat. Hist. Soc. 26:853-859.
Horin, O. 1989. Survey of White Stork
migration: Israel, spring 1984. Pp. 307-316 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status und Schutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Horin, O., and M. Adar. 1986. [Survey of the
White Stork migration, Israel, Spring 1984.]
Tzufit [Sunbird] 4:17-38, 138.
Horion, A. 1953. Koleopterologischer Beitrag zur
Kenntnis der Storchnahrung. Mitt. Bad.
Landesver. Naturk. NaturschutzFreib. Br. 6:7-16.
Hornberger, F. 1937. Neue Feststellungen ueber
Brutbestand, Ortstreu und Reife der
ostpreussischen Weiss-Storche im
Ermittlungskreis der Vogelwarte Rossitten.
Beitr. Fortpjlanzungsbiol Vogel 13:56-62.
Hornberger, F. 1938. Uberdie naturliche
Ragelung der ostpreussischen
Storchuebervolkerung durch Zugverluste und
Brutausfall im Jahre 1937. Beitr.
Fortplanzungsbiol. Vogel 14:168-175.
Hornberger, F. 1939. Zehn ostpreussische
Storch-Zahlugen. Orn. Monatsber. 47:166-170.
Hornberger, F. 1943a. Einige Ergebnisse
zehnjahriger Planarbeit in
'Storchforschungskreis Insterburg' der
Vogelwarte Rossitten./ Orn. 91:341-355.
Hornberger, F. 1943b. Ungewohnliche Storch-
Ankunftauch 1943. Vogelzug 14:109-112.
Hornberger, F. 1950a. Uber die Storche in
Baden und Wurttemberg und das
'Storungsjahr' 1949. Orn. 5^.47:98-108.
Hornberger, F. 1950b. Uber Storchzahlungen in
Deutschland und Osterreich. Orn. Beob.
47:108-126.
Hornberger, F. 1952. Ergebnisse der
Storchforschung. Naturwiss. Rundsch. 7:279-
284.
Hornberger, F. 1953. Vom Speisezettel des
Weissen Storches. Mitt. Bad. Landesver. Naturkd.
Naturschutz Freib. Br. 6:1-7.
Hornberger, F. 1954. Reifealter und Ansiedlung
beim Weissen Storch. Vogelwarte 17:114-149.
Hornberger, F. 1956a. Der Weissen Storch und
das Meer. Naturwiss. Monats. 63:182-185.
Hornberger, F. 1956b. Ist der Ruckgang des
Weissen Storches durch Kunstliche
Nestanlagen aufzuhalten? J/L Ver. Vaterl.
Naturk. Wurtt. 111:207-222.
Hornberger, F. 1956c. Das "Storchjahr" 1955 in
Baden-Wurttembergs. Mitt. Bad. Landesver.
Naturkd. Naturschutz Freib. Br. 6:330-338.
Hornberger, F. 1957a. Der Weisse Storch in
seinem Nahrungsrevier. Mitt. Ver. Naturwiss.
Math. £7/77*25:373^10.
Hornberger, F. 1957b. Der Weisse Storch in
Baden-Wurttemberg 1956. Beitr. Naturkd.
Forsch. Suedwestdtschl. Beih. 16:52-58.
Hornberger, F. 1958. Die WeiBstorche Baden-
Wurttembergs 1957. Mitt. Bad. Landesver.
Naturkd. Naturschutz Freib. Br. 7:197-204.
Hornberger, F. 1967. Der WeiBstorch. Neue
Brehm-Buechere Witt.lLuth. 375.
Hornberger, F., and G. Zink. 1960. Der Bestand
des WeiBstorchs in Baden-Wurttemberg 1958
und 1959. Jh. Ver. Vaterl. Naturkde. Wurttemberg
115:330-337.
Housse, P.R. 1945. Las Aves de Chile. University
of Chile, Santiago.
Houston, D.G. 1975. Ecological isolation of
African scavenging birds. Ardea 63:55-64.
Howard, G.W., and D.R. Aspinwall. 1984.
Aerial censuses of shoebills, saddle-billed
storks, and wattled cranes at the Bangweulu
Swamps and Kafue Flats, Zambia. Ostrich
55:207-212.
Howard, H. 1935. The Rancho La Brea Wood
Ibis. Com/or 37:251-253.
Howard, H. 1942. A review of the American
fossil storks. Carnegie Inst. Wash. Publ. 530:187-
203.
Howard, H. 1950. Fossil evidence of avian
evolution. Ibis 92:1-21.
Howard, H. 1962a. Fossil birds, with especial
reference to the birds of Rancho La Brea. Los
Angeles County Mus. Set. Ser. 17. 44pp.
Howard, H. 1962b. A comparison of avian
assemblages from individual pits at Rancho La
Brea, California. Los Ang. Cty. Mus. Contrib. Sci.
58:1-24.
Howard, H. 1969. Avian fossils from three
Pleistocene sites in central Mexico. Contr. Sci.
(Los Ang.) 172:1-11.
Howard, R.K., and K.W! Lowe. 1984. Predation
by birds as a factor influencing the
demography of an intertidal shrimp. J. Exp.
Mar. Biol. Ecol. 74:35-52.
Howell, A.H. 1932. Florida Bird Life. Coward-
McCann, New York.
Howell, J.C., Jr. 1930. Florida notes. Oologist
47:114-115.
Howell, J.C. 1941. Early nesting at Cape Sable,
Florida. Auk 58:105-106.
Howell, S.N.G., and B.M. de Monies. 1989.
Status of the Glossy Ibis in Mexico. Am. Birds
43:43-45.
Howells, W.W. 1968. Some interesting
Rhodesian nest records. Honeyguide 55:23-24.
Howes, J. 1985. Sarawak, East Malaysia.
Interwader Newsletter 5:4.
Howkes, J. 1985. Southward migration overview
1985, Sarawak, East Malaysia. Interwader
Newsletter 6.
Hoyt, R.D. 1906. Nesting of the Roseate
Spoonbill in Florida. Warbler 2:58-59.
Hu, H., and Y. Cui. 1990. The effects of habitat
destruction on the waterfowl of lakes in the
Yangtze and the Han river basins. Pp. 189-
193 in Matthews, G.V.T. (ed.), Managing
Waterfowl Populations, Proc. IWRB Symp.,
Astrakhan 1989. IWRB Special Publication
12. Slimbridge, U.K.
Hubaut, D. 1986. Aerial show of the Black Stork
(Ciconia nigra). Aves 23:184.
Hubbard, J.P., and Banks, R.C. 1970. The types
and taxa of Harold H. Bailey. Proc. Biol. Soc.
WwA. 83:321-332.
Hudec, K., D. Kondelka, and I. Novotny. 1966.
Ptactvo Slezska [Die Vogelwelt Schlesiens].
Herausgeber Slezske Muzeum Opava,
Tschechoslowakei.
Hudson, W.H. 1920. Birds of La Plata, Vol. 2.
J.M. Dent, London.
Hudson, W.H. 1951. Letter on the Ornithology of
Buenos Aires. Cornell University Press, Ithaca.
93pp.
Hue, F. 1953. Note sur 1'Ibis noir Eudocimus
eremita (L.) au Maroc. Alauda 11:194.
Hughes, R., and A. Owen. 1988. Hand-rearing
the Scarlet Ibis Eudocimus ruber. Avic. Mag.
94:96-100.
Hughes, R.A. 1970. Notes on the birds of the
Mollendo District, southwest Peru. Ibis
112:229-241.
Hughes, R.A. 1984. Further notes on Puna bird
species on the coast of Peru. Condor 86:93.
Hume, A.O. 1888. The birds of Manipur,
Assam, Sylhet and Cachar. Stray Feathers 11:1—
353.
Hume, A.O., and E.W. Gates. 1890. The Nests
and Eggs of Indian Birds, 2nd edn. R.H. Porter,
London.
Hummel, D. 1975. Zum Wegzug des Weiss-
Storchs (Ciconia ciconia) im Israelischen
Kuestengebiet. Vogelwarte 28(2): 131-134.
Humphrey, C. 1963. Roseate spoonbills at New
Smyrna Beach. Fla. Nat. 36:27.
Humphrey, P.S., D. Bridge, P.W. Reynolds, and
R.T. Peterson. 1970. Birds of Isia Grande (Tierra
del Fuego). Smithsonian, Washington. 411pp.
Humphrey, R.C. 1987. Range expansion and
new breeding record for the Glossy Ibis in
Massachusetts. Bird Obs. (Mass.) 15:173-177.
Hundertmark, C.A. 1974. Breeding range
extensions of certain birds in New Mexico.
Wilson Bull. 86:298-300.
Hunter, R. 1978. White Ibis in Osage County,
Oklahoma. Bull. Okla. Orn. Soc. 11(3):20.
Huntley, B.J., and M.A. Huntley. 1974. Hadada
Ibis (Bostrychia hagedash) new distributional
data. Ostrich 45:133.
Hussain, Z. 1979. Birds of Bangladesh.
Department of Films and Publications,
Government of the People's Republic of
Bangladesh, Dacca.
Hutson, H.W. 1944. Observations of the White
Stork 1942/43. Bull. Zool. Soc. Egypt 10(suppl.
6):6-8.
Hutton, R.F. 1964. A second list of parasites
from marine and coastal animals of Florida.
Trans. Am. Microsc. Soc. 83:439-447.
Hutton, R.F., and F. Sogandares-Bernal. 1960. A
list of parasites from marine and coastal
animals of Florida. Trans. Am. Microsc. Soc.
79:287-292.
Huxley, J.S. 1960. The openbill's open bill: a
teleonomic enquiry. Zool.Jahrb. Abt. Syst. Oekol.
Geogr. Tiere 88:9-30.
Huxley, J. 1962. The Openbill Stork Anastomus
oscitans. Ibis \M\\\2.
Huxley, T.H. 1867. On the classification of birds;
and on the taxonomic value of the
modifications of certain of the cranial bones
observable in that class. Zool. Soc. Lond. Proc.
1867:415-472.
Hyem, E.L. 1957. Notes on a trip to the
Macquarie Marshes. Emu 57:6—8.
Hyrtl, J. 1864. Neue Wundernetze und Geflechte
bei Voegeln und Saugethieren. Denkschr.
Kaiserlichen Akad. Wissensch., Wien, Math.-
Naturwiss. Classe 22:113-152.
IUCN (in press) The IUCN Directory of
Afrotropical Protected Areas. IUCN Monitoring
Centre and Commission on National Parks
and Protected Areas, Cambridge, and Gland,
Switzerland.
Ihering, H. v., and R. v. Ihering. 1907. Catalogos
da Fauna Brazileira, vol. 1. La Aves do Brazil.
Ilyinskiy, I.V., Yu. B. Pukinskiy, and S.A.
Fetisov. 1988. [Acoustic communication by
Black Stork (Ciconia nigra} chicks.] Tr. Vsesoyuz
Orn. Ofcfaft. 2:140-151.
Imboden, C., and D. Parish. 1986. The Milky
Stork recovery action plan. Unpublished
project proposal. ICBP. Cambridge, England.
Imhof, T.A. 1962. Alabama Birds. University of
Alabama Press, University. 591pp.
Inglis, C.M. 1898. Note on the nidification of the
White-necked Stork (Dissoura episcopus). J.
Bombay Nat. Hist. Soc. 12:217-218.
Inglis, C.M., W.L. Travers, H.V. O'Donel, and
E.O. Shebbeare. 1920. A tentative list of the
vertebrates of the Jalpaiguri District, Bengal,
part 3. J. Bombay Nat. Hist. Soc. 27:151-162.
Ingrand, A. 1959. Cature d'un Ibis Falcinelle en
Vendee. Alauda 27:69.
Inkapatanakul, W. 1986. Factors influencing the
survival of the Asian Open-billed Stork
fledgling Anastomus oscitans (Boddaert) at Wat
Phai Lorn and Wat Umpuwaram Non-hunting
Area, Pathum Thani Province (Thailand).
350
M.S. Thesis. Kasetsart University, Bangkok,
Thailand. 67pp.
Inskipp, C., and T. Inskipp. 1985. A Guide to the
Birds of Nepal. Groom Helm, London.
International Council for Bird Preservation.
1966. Survival service commission red data
book. ICBP, Cambridge.
International Council for Bird Preservation.
1985. Council of Europe Important Bird
Inventory. Unpublished report. ICBP,
Cambridge.
International Union for Conservation of Nature
and Natural Resources. 1966. Japanese
Crested Ibis. Survival Service Commission Red
Data Book. Merges, Switzerland.
International Union for the Conservation of
Nature and Natural Resources. 1990. Directory
of Wetlands of International Importance. Ramsar
Convention Bureau, Gland, Switzerland.
796pp.
lordachescu-Lazarescu, D. 1965. New data
concerning host-parasite relationships
observed in parasitic infections in Ciconia nigra
L. Trav. Mus. Hist. Nat. 'Grigore Antipa'5:481-
485.
Iredale, T. 1956. Birds of New Guinea, vol. 1.
Georgian House, Melbourne.
Irmscher, S., and R. Irmscher. 1979. White Ibis
at Centerfield, Kentucky. Kentucky Warbler
55:34-35.
Irwin, M.P.S. 1980. A misplaced Marabou
breeding locality. Honeyguide 102:36.
Irwin, M.P.S. 1981. The Birds of Zimbabwe. Quest
Publishing, Salisbury.
Ishii, S., A. Hattori, H. Sakai, S. Wakabayashi,
and M. Wada. 1984. Annual changes in
plasma gonadotropins and sex steroid in the
White Ibis in captivity. Ann. Meet. Zool. Soc.
Japan 55:974.
Iskander, J. 1985. Survey of waders, herons,
egrets, storks, and terns in southeastern coast
of Sumatra, Indonesia. Proc. East-Asian Bird
Protect. Con/. 3:108-117.
Itoh, S. 1986. Records of the Oriental Ibis
Threskiornis melanocephalus in Japan. Tori
34:127-143.
Ivey, G.L., and D.J. Severson. 1984. White-faced
Ibis (Plegadis chihi) nesting in the San Joaquin
Valley of California U.S.A. Condor 86:492-493.
Ivey, G.L., M.A. Stern, and C.G. Carey. 1988.
An increasing White-faced Ibis population in
Oregon. West. Birds 19:105-108.
Jaarsveld, J. v. 1979. The Bald Ibis. Fauna Flora
(Transv.) 35:12-13.
Jaarsveld, J. v. 1980. Bald Ibis—a master of the
air. Afr. Wildl. 34(6):20-23.
Jackson, FJ. 1938. The Birds of Kenya Colony and
the Uganda Protectorate, vol. 1. Gurney and
Jackson, London. 542pp.
Jackson, P., (ed.). 1978. Turkey Project 1062
Bald Ibis. WWF Yb. 1977-1978.
Jacob, J., and D.E. Pomeroy. 1979. The feather
lipids of the Marabou Stork (Leptoptilos
crumeniferus). Comp. Biochem. Physiol. 64B:301-
303.
Jacobaeus, O. 1677-79. Anatome Ciconiae. Acta
Med. Phil. Haf., 5:247-249. Collect. Acad., 1757,
4:356-357.
Jacot-Guillarmod, C. 1936. Catalogue of the
birds of Basutoland. S. Afr. Avifauna 8.
Jacot-Guillarmod, C. 1965. The Openbill
Anastomus lamelligerus in the Eastern Cape
Province. Ostrich 36:138.
Jaensch, R.P., and C. Auricht. 1985. Another
breeding colony of the Glossy Ibis in South
Australia. S. Aust. Orn. 29:208.
Jagans, M., and A. Stipniece. 1989. 50 years
(1934-1984) population trends of the White
Stork in Latvia. Pp. 145-152 in Rheinwald, G.,
J. Ogden, and H. Schulz (eds), Weiflstorch:
Status undSchutz, Proc. 1st Int. Stork Conserv.
 Symp., Walsrode, 1985, Schriftenr. DBA,
Braunschweig, Germany.
Jahn, H. 1942. Zur Oekologie und Biologic der
Vogel Japans. J. Om. 90:4-302.
Jakab, B. 1987. Breeding numbers of the stork
(C. ciconia) in Hungary, 1958-1984. Falke
34:47-50.
Jakab, B. 1989. Der WeiBstorch in Ungarn:
Bestandsveranderungen und ihre Ursachen.
Pp. 105-114 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weifistorch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
James, C., N. N-Gyan, and G. Hislop. 1984.
ICBP/IWRB neotropical wetlands report. National
Report, Trinidad and Tobago. Cambridge,
England.
Janez, G. 1987. Black Stork (C. nigra) breeding at
Ljubljiana Morass. Acrocephalus 33:37-39.
Janson, V.S. 1964. Wood Ibis sighting near
Lansing. Jack-Pine Warbler 42:229.
Janssen, R.B. 1967. White-faced ibis at Heron
Lake. loon 39:108.
Jarvis, C. (ed.). 1966. Species of animals bred in
zoos and aquaria during 1964. Int. Zoo Yb.
6:385-431.
Jasmin, A.M., D.E. Cooperrider, C.P. Powell,
andJ.N. Baucom. 1972. Enterotoxemia of
wildfowl due to Cl. perfringens Type C. J. Wildl.
Dis. 8:79-84.
Jehl, J.R., Jr. 1966. Fossil birds from the Sand
Draw local fauna (Aftonian) of Brown County,
Nebraska. Auk 83:669-670.
Jencks, F.T. 1884. The Roseate Spoonbill in
Florida rookeries. Random Notes on Nat. Hist.
1(3):4; 1(4):7; l(5):4-5; 1(6):4.
Jenkins, C. 1968. White Ibis near Perth. West.
Aust.Nat. 11:46.
Jensen, J.R., H.E. Mackey, and M. Coulter.
1986. Remote sensing of endangered species
foraging habitats: a Wood Stork example Pp.
485-494 in Photogrammetry. Proceedings of the 1986
Joint Meeting of the American Congress on Surveying
and Mapping and the American Society for
Photogrammetry and Remote Sensing. Falls Church,
Virginia.
Jensen, J.R., M.E. Hodgson, M.C. Coulter, and
H.E. Mackey, Jr. 1989. Feasibility study of
Wood Stork foraging habitat mapping using
Landsat multispectral data. Pp. 1059-1068 in
Sharitz, R.R., and J.W. Gibbons (eds),
Freshwater Wetlands and Wildlife. D.O.E.
Symposium Series No. 61. U.S.D.O.E., Oak
Ridge, Tenn.
Jensen, J.V., andj. Kirkeby. 1980. The Birds of
The Gambia. Aros Nature Guide, Aathus,
Denmark.
Jerdon, T.C. 1864. The Birds of India, vol. III.
Wyman and Co., Calcutta.
Jernelov, A., H. Kossmann, and P.-E. Larsson.
1984. Storkprojektet i Aneboda. Fauna Flora
(Stockh.) 79:1 13-1 16.
Jerrentrup, H. 1989. Vergleich zweier Teil-
Populationen des WeiBstorchs (Ciconia ciconia)
im Nestos-Delta, Nordost-Griechenland. Pp.
127-136 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiftstorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Jespersen, P. 1951. Contribution a la
connaisance de la migration des Cicognes
blanches (Ciconia ciconia(L.)) en Afrique du
Nord. Proc. Int. Orn. Congr. 10:344-350.
Jewell, S.D. 1987 (1989). Larval dermestid
infestation of nestling Wood Storks (Myctena
americana) in Georgia. Oriole 52:1 1-13.
Jez, M. 1987. The White Stork (Ciconia ciconia L.)
in Slovenia in the year 1979. Varstvo Narave
[Nat. Conserv.] 13:79-92.
Ji, J., and X. Yu. 1990. [The numerical
distribution of storks and cranes in Shandong
Province.] Zoo/. Res. 11:46.
Jia, D. et al. 1990. Histological research on
digestive organs of Oriental White Storks.
Chinese Wildlife 1990(6):33-35.
Jiang F. 1991. Preliminary observations on
captive breeding of Oriental White Storks. Pp.
213-218 in Coulter, M.C., Q. Wang, and C.S.
Luthin (eds), Biology and Conservation of the
Oriental White Stork (Ciconia boyciana).
Savannah River Ecology Laboratory, Aiken,
South Carolina, USA.
Jobanek, G.A. 1987. Early records of the White-
faced Ibis in Oregon. Oregon Birds 13:210-215.
Jobert, C. 1873. Recherches sur la structure
intime du bee de la Spatule (Platalea). C. R.
Acad. Sci. Paris 75:1780-1782.
Johansen, H. 1954. Uberden turkestanischen
WeiBstorch (Ciconia c. asiatica). Vogelwarte
17:162.
Johansen, H. 1958. Die Vogelfauna
Westsibiriens. J. Orn. 99:67-80.
Johansen, H., and A. Bjerring. 1955. Bestanden
af Stork (Ciconia ciconia(L.)) i Danmark 1952-
1954. Dan. Orn. Foren. Tidsshr. 49:114-125.
Johansen, H., and A. Bjerring. 1962. Bestanden
af Stork (Ciconia ciconia(L.)) i Danmark 1955-
1960. Dan. Orn. Foren. Tidsshr. 56:40-55.
Johns, F.L., and Erickson, J.E. 1964. White Ibis
observations in North Carolina. Chat 28:94-
95.
Johnson, A.W. 1965. The Birds of Chile and
Adjacent Regions of Argentina, Bolivia and Peru,
vol. I. Platt Establecimientos Graficos, Buenos
Aires, Argentina.
Johnson, J.M. 1984. A large flock of migrating
White Storks Ciconia ciconia. J. Bombay Nat.
Hist. Soc. 81:466-467.
Johnson, K.C. 1934. Notes on the nesting birds
of the Orange Free State. Ostrich 5:49-56.
Johnson, L.R. 1958. Field notes on some of the
birds of Iraq. Iraq Nat. Hist. Mus. Publ. 16:1-
32.
Johnson, M., V. Nagulu, and J.V. Ramana Rao.
1990. Some observations on the feeding
ecology of the Openbill Stork at Kolleru
wetland habitat in Andhra Pradesh. Pp. 49-50
in Seminar on Wetland Ecology and Management.
Bombay Nat. Hist. Soc., Bharatpur, India.
Johnson, R. 1990. An, appreciation of storks. Zoo
Life 1:38-44.
Johnson, R., and C.S. Luthin. 1990.
Ciconiidae—current world report and use of
Yellow-billed Stork (Mycteria ibis) as a model
species for captive management of endangered
storks. Pp. 93-104 in Dresser, B.L. et al. (eds),
Proceedings 5th World Conference on Breeding
Endangered Species in Captivity, Cincinnati, Ohio.
Johnson, R.E., M.C. Coulter, C.S. Luthin, C.E.
King, and A.J. Valenzuela. 1987a. Storks:
status, conservation, and captive breeding.
Colonial Waterbirds 10:236-241.
Johnson, R.E., M.C. Coulter, C.S. Luthin, C.E.
King, and A.J; Valetizuela. 1987b. Storks:
status, conservation, and captive breeding. AAZPA
Annu. Proc., pp. 279-296.
Johnston, D.W., C.H. Blake, and D.W. Buden.
1971. Avifauna of the Cayman Islands. Q.J.
Fla. Acad. Sci. 34(2): 141-156.
Johnston, H. 1902. On the occurrence of
Balaeniceps rex on Lake Victoria. Ibis (ser. 8).
2:334-336.
Johnston, J.W., and K.L. Bildstein. 1990.
Dietary salt as a physiological constraint in
White Ibis breeding in an estuary. Physiol.
Zoo/. 63:190-207.
Jolly, A. 1980. A World Like Ours-Man and Nature
in Madagascar. Yale University Press, New
Haven.
Jones, P. 1989. Dwarf Olive Ibis found on Sao
Tome. Spec. Group Storks, Ibis, Spoonbills
Newsletter 2:9-10.
351
Jones, PJ. 1988. Ibises in the 'OBO'. World
Birdwatch 10:5.
Jong, B.HJ. d. 1981. Voorkomen en talrijkheid
van een aantal soorten watervogels langs de
kust van de Guiana's in 1981. Unpublished
report, Dienst's Lands Bosbeheer/Stichting.
Jong, B.HJ. d. 1982. Voorkomen en talrijkheid
van een aantal soorten watervogels langs de
kust van Suriname en Frans-Guyana in juni
1982. Unpublished report, Dienst's Lands
Bosbeheer/STINASU, Paramaribo.
Jong, B.HJ. d. 1983. Inventarisatie Rode Ibis
langs de kust van de Guianas in 1982. Vogeljaar
31:156-157.
Jong, B.HJ. d., and A.L. Spaans. 1980.
Voorkomen en talrijkheid van een aantal
soorten watervogels langs de kust van
Suriname in juni 1980. Unpublished report,
Dienst's Lands Bosbeheer, Paramaribo/
Rijksinstituut voor Natuurbeheer, Arnhem.
Jong, B.HJ. d., A.L. Spaans, and M. Held.
1984. Waterfowl and wetlands in Suriname.
Contribution to the IWRB/ICBP Neotropical
Wetlands Project. RIN Contributions to
Research on Management of Natural
Resources No. 1984-1.
Jonkers, D.A. 1975. De stand van de Ooievaar in
1975. Het Vogeljaar 23:261-266.
Jonkers, D.A. 1980. Op het spoor van de
ooievaar. Lepelaar 68:92—95.
Jonkers, D.A. 1987a. Foerageergebieden en voedsel
van de Ooievaars in Schoonerwoerd.
Ooievaarswerkgroep Schoonerwoerd, Vianen.
Jonkers, D.A. 1987b. Hoe het in 1987 ging met
de ooievaars in Nederland. Vogeljaar 35:328-331
Jonkers, D.A. 1988. Lotgevallen van ooievaars in
Nederland in 1988. Fo£*/>zr36:241-243.
Jonkers, D.A. 1989. Status and conservation of
the White Stork in the Netherlands: a review.
Pp. 45-54 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiftstorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Jonkers, D.A. 1990a. Nederland en zijn ooievaars
in 1989. Vogeljaar 38:15-18.
Jonkers, D.A. 1990b. Broedende en trekkende
Ooivaars in 1990 in Nederland. Vogeljaar
39:252-254.
Jergensen, H.I., and C.I. Blackburne. 1941.
Glossarium Europae Avium. Munksgaard,
Kobenhavn.
Jourdain, F.C.R. 1922. Les oiseaux de la foret de
Marmara et des environs de Rabat. Rev. Fr.
Orn. 7:149-153.
Jourdain, F.C.R. 1934. The Bald Ibis (Comatibis
eremita). Ool. Rec. 14(1): 1-5.
Jovetic, R. 1960a. White Stork, Ciconia ciconia, in
Macedonia. Larus 14:75-83.
Jozefik, M. 1960b. Legi podwojne u ibisow
kasztanowatych, Plegadis falcinellusfalcinellus
(L.) w delcie Dniestru. [The double broods of
the glossy ibises, Plegadis falcinellus falcinellus
(L.) in the Dniester delta.] Acta Orn. (Warsaw)
5:367-377.
Jovetic, R. 1961. The life of the White Stork,
Ciconia ciconia^ in Macedonia. Larus 15:28-99.
Ju, C. et al. 1990 Oriental White Storks and their
preservation strategies in Jilin Province.
Chinese Wildl. 1990(3):9-11.
Jubb, R.A. 1952. Status of the White Stork,
Ciconia ciconia, in Southern Rhodesia. Ostrich
23:125-126.
Junge, G.C.A., and G.F. Mees. 1958. The
avifauna of Trinidad and Tobago. Zoo/. Verh.
(Leiden) 37:1-172.
Junge, G.C.A., and J. Taapken. 1953. Resultaten
van het ringonderzoek betreffende de
vogeltrek, ingesteld door het Rijksmuseum van
Natuurlijke Historic te Leiden, XXXIX
(1952), 1. Limosa 26:80-100.
Juran, V. 1955. White Stork's strange nest-
building. Aquila 59-62:434-435.
Kaatz, C. 1982. Sicherung von WeiBstorchhorsten
auf Schornsteinen. Falke 29:127-130.
Kaatz, C., and K. Spange. 1985. Aufzucht eines
abwurfgefahrdeten Jungstorches. Falke 32:45-
47.
Kahl, M.P. 1962. Bioenergetics of growth in
nestling Wood Storks. Co^or 64:169-183.
Kahl, M.P. 1963a. Technique for sampling
population density of small shallow-water fish.
LimnoL & Oceanogr. 8:302-304.
Kahl, M.P. 1963b. Thermoregulation in the
Wood Stork, with special reference to the role
of the legs. PhysioL Zool. 36:141-151.
Kahl, M.P. 1964. Food ecology of the Wood
Stork (Mycteria americana) in Florida. Ecol.
Monogr. 34:97-117.
Kahl, M.P. 1965. Whale-headed Stork-a
feathered riddle. Africana 2:19-20.
Kahl, M.P. 1966a. Comparative ethology of the
Ciconiidae. Part 1. The Marabou Stork,
Leptoptilos crumeniferus (Lesson). Behaviour
27:76-106.
Kahl, M.P. 1966b. A contribution to the ecology
and reproductive biology of the Marabou
Stork (Leptoptilos crumeniferus) in East Africa. J.
Zool. (Land.) 148:289-311.
Kahl, M.P. 1967a. Behavioural reactions to
hyperthermia in Scopus umbrella and Balaeniceps
rex. Ostrich 38:27-30.
Kahl, M.P. 1967b. Observations on the
behaviour of the Hamerkop, Scopus umbrella, in
Uganda. Ibis 109:25-32.
Kahl, M.P. 1968. Recent breeding records of
storks in East Africa. J. East Afr. Nat. Hist. Soc.
Natl. Mus. 27:67-72.
Kahl, M.P. 1970 [1971]. Observations on the
breeding of storks in India and Ceylon. J.
Bombay Nat. Hist. Soc. 67:453-461.
Kahl, M.P. 1971a. The courtship of storks. Nat.
Hist. 80:36-45.
Kahl, M.P. 1971b. Flapping rates of storks in
level flight. Auk 88:428.
Kahl, M.P. 1971c. Food and feeding behavior of
Openbill Storks. J. Orn. 112:21-35.
Kahl, M.P. 197Id. Observations on the breeding
of the Abdim's Stork at Lake Shala, Ethiopia.
Ostrich 42:233-241.
Kahl, M.P. 197le. Observations on the Jabiru
and Maguari Storks in Argentina, 1969. Condor
73:220-229.
Kahl, M.P. 1971f. Social behavior and
taxonomic relationships of storks. Living Bird
10:151-170.
Kahl, M.P. 1971g. Spread-wing postures and
their possible functions in the Ciconiidae. Auk
88:715-722.
Kahl, M.P. 1972a. Comparative ethology of the
Ciconiidae. Part 2. The Adjutant Storks,
Leptoptilos dubius (Gmelin) and L.javanicus
(Horsfield). Ardea 60:97-111.
Kahl, M.P. 1972b. Comparative ethology of the
Ciconiidae. Part 3. The Wood Storks (genera
Mycteria and Ibis). Ibis 114:15-29.
Kahl, M.P. 1972c. Comparative ethology of the
Ciconiidae. Part 4. The 'Typical' Storks
(genera Ciconia, Sphenorhynchus, Dissoura, and
Euxenura). Z. Tierpsychol. 30:225-252.
Kahl, M.P. 1972d. Comparative ethology of the
Ciconiidae. Part 5. The Openbill Storks (genus
Anastomus).J. Orn. 113:121-137.
Kahl, M.P. 1972e. A revision of the Family
Ciconiidae (Aves). J. Zool. (Lond.) 167:451-
461.
Kahl, M.P. 1972f. Thestork: a taste for survival.
Pp.254—269 in The Marvels of Animal Behavior.
National Geographic Society. Washington,
D.C.
Kahl, M.P. 1973a. Comparative behavior and
ecology of African storks. Pp. 137-141 in
National Geographic Society Research Reports, 1966
projects. National Geographic Society,
Washington DC.
Kahl, M.P. 1973b. Comparative ethology of the
Ciconiidae. Part 6. The Blacknecked,
Saddlebill and Jabiru storks (genera
Xenorhynchus, Ephippiorhynchus and Jabiru).
Condor 75:17-27.
Kahl, M.P. 1974. Comparative behavior and
ecology of Asian storks. Pp. 173-176 in
National Geographic Society Research Reports, 1967
projects. National Geographic Society,
Washington DC.
Kahl, M.P. 1976. Comparative behavior and
ecology of American and European storks. Pp.
173-176 in National Geographic Society Research
Reports, 1968projects. National Geographic
Society, Washington DC.
Kahl, M.P. 1978. Wonders of Storks. Dodd, Mead
Publishers, New York. 94pp.
Kahl, M.P. 1979a. Family Balaenicipitidae. Pp.
252-253 in Mayr, E., and G.W. Cottrell (eds),
Check-list of Birds of the World, vol. 1, 2nd edn.
Museum of Comparative Zoology, Cambridge,
Massachusetts.
Kahl, M.P. 1979b. Family Ciconiidae. Pp. 245-
252 in Mayr, E., and G.W. Cottrell (eds),
Checklist of Birds of the World, vol. 1, 2nd edn.
Museum of Comparative Zoology, Cambridge,
Massachusetts.
Kahl, M.P. 1979c. Suborder Scopi; Family
Scopidae. Pp. 244-245 in Mayr, E., and G.W.
Cottrell (eds), Check-list of Birds of the World,
vol. 1, 2nd edn. Museum of Comparative
Zoology, Cambridge, Massachusetts.
Kahl, M.P. 1981. Welt der Stoerche. Parey,
Hamburg.
Kahl, M.P. 1983. Breeding displays of the
African Spoonbill Platalea alba. Ibis 125:324-
338.
Kahl, M.P. 1987a. An overview of the storks of
the world. Colonial Waterbirds 10:131-134.
Kahl, M.P. 1987b. A portfolio of storks of the
world. Colonial Waterbirds 10:135-150.
Kahl, M.P. 1987c. The Royal Spoonbill. Natl.
Geogr. Mag. 171:280-284.
Kahl, M.P. 1988a. Breeding displays of
Australian spoonbills. Natl Geogr. Res. 4:88-
111.
Kahl, M.P. 1988b. The stork brought him. Int.
Wildl. 18:44-50.
Kahl, M.P. and LJ. Peacock. 1963. The bill-
snap reflex: a feeding mechanism in the
American Wood Stork. Nature 199:505-506.
Kahl, M.P., and E. Schiiz. 1972. Zur Benennung
und zur taxonomischen Gruppierung der 17
Arten Storche (Ciconiidae). Vogelwarte 26:277-
280.
Kale, H.W., II. 1965. Nestling predation by
herons in a Georgia heronry. Oriole 30:69-70.
Kaleta, E.F., and N. Kumerfeld. 1983.
Herpesviruses and Newcastle disease viruses
in White Storks (Ciconia ciconia). Avian Pathol.
12:347-352.
Kaleta, E.F., T. Mikami, H.-J. Marschall, U.
Heffels, M. Heidenreich, and B. Stiburek.
1980. A new herpesvirus isolated from Black
Storks (Ciconia nigra). Avian Pathol. 9:301-310.
Kalitsch, L. 1940. Sechser- und Siebenergelge
beim Weissen Storch. Beitr. Fortpflanzungbiol.
Vogell6:W6.
Kamel, I. 1971. Hybrid between Japanese Stork
and European White Stork. Anim. Zoos 23(11)
(no.262):370.
Kaneko, K., and T. Kadosaka. 1977. Notes on
three species of biting lice (Mallophaga) found
on Japanese White Stork (Ciconia c. boyciana) in
Japan. Misc. Rep. Yamashina Inst. Orn. 9:29-33.
Kania, W. 1985. [Results of bird ringing in
Poland. Migrations of White Stork Ciconia
ciconia.] Ada Orn. (Warsaw) 21:1-41.
Kania, W. 1988. Investigations of White Stork
Ciconia ciconia hatching phenology based on bill
352
measurements of nestlings. Ring Int. Orn. Bull.
12:13-19.
Kanyamibwa, S., A. Schierer, R. Pradel, and
J.D. Lebreton. 1990. Changes in adult annual
survival rates in a western European
population of the White Stork Ciconia ciconia.
Ibis 132:27-35.
Kappler, A. 1854. Zesjaren in Suriname.
Dannenfelser, Utrecht.
Kappler, A. 1881. Hollandisch- Guiana.
Commissions-Verlag von W.Kohlhammer,
Stuttgart.
Kappler, A. 1887. Surinam, sein Land, seine Natur,
Bevolkerung und seine Kultur-verhaltnisse.
Cotta'sche Verlagsbuchhandlung, Stuttgart.
Kasoma, P.M.B., and D.E. Pomeroy. 1987. The
status and ecology of storks and the Shoebill in
East Africa. Colonial Waterbirds 10:221-228.
Kasparak, M., and A. Kilic. 1989a. Zum Zug
des WeiBstorchs (Ciconia ciconia) durch die
Turkei. Pp. 297-306 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weifstorch: Status
undSchutz, Proc. 1st Int. Stork Conserv. Symp.,
Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Kasparek, M., and A. Kilic. 1989b.
Brutverbreitung und Bestandsentwicklung des
WeiBstorchs in der Turkei. Pp. 161-174 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weifstorch: Status undSchutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Katz, Y., V. Liepa, and J. Viksne. 1988.
Orientation research in the Latvian S.S.R. in
1982-1985. Proc. Int. Orn. Congr. 19:1919-1931.
Keil, W., and R. Rossbach. 1969.
Bestandsveranderungen des WeiBstorchs—
Ciconia ciconia—in Hessen von 1948 bis 1968.
Luscinia 40:230-249.
Keimer, L. 1954-1955. Interpretation de
plusieurs representations anciennes d'ibis.
Chron. d'Egypt (Brussels) 29:237-250, 30:46
(addendum).
Keimer, L. 1955. The Egyptian Ibis. Egypt Travel
Mag. 16:16-19.
Keith, G.S., A.D. Forbes-Watson, and D.A.
Turner. 1974. The Madagascar Crested Ibis, a
threatened species in an endemic and
endangered avifauna. Wilson Bull. 86:197-199.
Keith, S. 1968. Notes on birds of East Africa
including additions to the avifauna. Amer. Mus.
Novit. 2321:1-15.
Keller, C.E. 1966. Status of the Ciconiiformes in
Indiana. Indiana Audubon Q. 44:56-86.
Keller, C.E., J.A. Spendelow, and R.D. Greer.
1984. Atlas of wading bird and sea bird nesting
colonies in coastal Louisiana, Mississippi, and
Alabama 1983. U.S. Fish Wildl. Serv. Biol. Rep.
FWS/OBS-84/13. US Dept of the Interior,
Washington DC.
Kemp, A.C. 1974. The distribution and status of
the birds of the Kruger National Park. Koedoe,
Monogr. 2:1-352.
Kemp, A.C. 1980. The importance of the Kruger
National Park for bird conservation in the
Republic of South Africa. Koedoe 23:99-122.
Kemp, A.C., and M.I. Kemp. 1976. Random
notes on some Sarawak birds. Sarawak Mus.J.
24:273-276.
Kemper, J.H. 1985. Foraging strategy in the
Spoonbill (Platalea leucorodia). Verslagen en
Technische Gegevens, Instituut voor
Taxonomische Zooelogie, Universiteit van
Amsterdam, No. 44.
Kemper, J. 1988. Zalm voor de Lepelaars. Vogels
46:147-148.
Kemper, J. 1989. [Fishway for sticklebacks to
save the Spoonbill (Platalea leucorodia) in
Noord-Holland.] Levende Nat. 90:2-6.
Kennard, F.H. 1915. The Okaloacoochee
Slough. 4M* 32:154-166.
Kennerley, P.R. 1987. A survey of the birds of
 the Poyang Lake Nature Reserve, Jiangxi
Province, China, 29 December 1985—4
January 1986. Hong Kong Bird Rep. 1984/
1985:97-111.
Kennerley, P.R. 1990. A review of the status and
distribution of Black-faced Spoonbill. Hong
Kong Bird Rep. 1989:116-125.
Kerautret, L. 1967. Observations
ornithologiques dans le Nord de la Grande-
Kabylie (Algerie) (Mars 1961-Aout 1962).
Oiseau Rev. Fr. Orn. 37:221-239.
Kerr, J.G. 1901. On the birds observed during a
second zoological expedition to the Gran
Chaco. Ibis (ser. 8) 1:215-236.
Kesteloot, E. 1989. A review of the history and
recent status of the White Stork in Belgium.
Pp. 43^4 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weifistorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DBA, Braunschweig,
Germany.
Keve, A. 1967. Uber die Arealveranderungen des
Sichlers Plegadisfalcinellus. Zoo/. Abh. (Dres.)
29:159-175.
Keve, A., and E. Patkai. 1959. Hungarian ringed
birds in Africa. Ostrich Suppl. 3:321-330.
Khacher, L J. 1976a. Some notes on the White
Stork Ciconia ciconia, the Black Drongo dicrurus
adsimilis and the Starling Sturnus vulgaris. J.
Bombay Nat. Hist. Soc. 73:389-390.
Khacher, S. 1976b. Occurence of the Black Stork
(Ciconia nigra) in Saurashtra. J. Bombay Nat.
Hist. Soc. 73:390-391.
Khan, M.A.R. 1982. Wildlife of Bangladesh, a
Checklist. University of Dhaka, Dhaka.
Khan, M.A.R. 1984. Conservation of storks and
ibises in Bangladesh. Tigerpaper ll(4):2-4.
Khan, M.A.R. 1987. Conservation of storks and
other waterbirds in Bangladesh. Colonial
Waterbirds 10:229-235.
Khan, M.A.R. 1989. Migratory status of the
White Stork in the United Arab Emirates. Pp.
325-236 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weifistorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Khan, R. 1987. Stork story. Horizons 2:19-21.
Kidd, E. 1978. Some notes on the birds of
Brunei. Brunei Mus.J. 4:115-164.
Kidiralev, V.V. 1977. [Mountain Goose,
European White and Black Storks in the Tyan
Shan Mountain Range in Kirgiziya.] Pp. 225-
226 in Winter, S. (ed.), VII All-Soviet
Ornithological Conference. Zoological Institute of
the Academy of Science of the Ukranian SSR,
Kiev.
Kieser, G.A., and J.A. Kieser. 1977. Spoonbills
breeding in the northern Karoo. Bokmakierie
29:53.
Kieser, J.A., and G.A. Kieser. 1978. Birds of the
De Aar District. South. Birds 5:1-46.
Kilham, L. 1980. Association of Great Egret and
the White Ibis. J. Field Orn. 51:73-74.
Kilham, L. 1984. American Crows (Corvus
brachyrhynchos) robbing Great Egrets
(Casmerodius alsubs) and White Ibis (Eudocimus
albus) of large, eel-like salamanders. Colonial
Waterbirds 7:143-145.
Killerman, S. 1911. Der Waldrapp Gesners neue
Zeugnisse fur sein ehemaliges Vorkommen in
Mitteleuropa. Zoo/. Ann. Wurburg^Tl.
King, B.F., and E.G. Dickinson 1975. A Field
Guide to the Birds of South-East Asia. Collins,
London.
King, C.E. 1988. An ethological comparison of
three storks: Ciconia boyciana, C. ciconia, and C.
maguari. M.S. Thesis, Oklahoma State
University, Stillwater, Oklahoma.
King, C. 1990. Reproductive management of the
Oriental White Stork Ciconia boyciana in
captivity. Int. Zoo. Yb. 29:85-90.
King, C., and K. Brouwer. 1991. Conservation
and captive management of Indian storks.
Zoos' Print 6(l):\-3.
King, C.E., and M.C. Coulter. 1989. Status of
storks in zoos: 1987 survey. Int. Zoo Yb.
28:225-229.
King, C.E., and S. Patzwahl. 1987. First
successful hatching of the Eastern White Stork
outside of China. Int. Zoo News 34(4): 17-18.
King, K.A., E.L. Flickinger, and H.H.
Hildebrand. 1978. Shell thinning and pesticide
residues in Texas aquatic bird eggs. Pest.
Monit.]. 12:16-21.
King, K.A., D.L. Meeker, and D.M. Swineford.
1980. White-faced Ibis populations and
pollutants in Texas, 1969-1976. Southwest Nat.
25:225-239.
King, W.B. 1978. Red Data Book. Vol. II. Aves.
Int. Union Conserv. Nat., Morges,
Switzerland.
King, W.B. 1981. Endangered Species of the World.
The ICBP Bird Red Databook. Smithsonian
Institution Press, Washington, D.C.
Kinzelbach, R.K. 1986. New records of Goliath
Heron, Yellow-billed Stork, Blyth's Reed
Warbler, and Clamorous Reed Warbler in the
Middle East. Bull. Orn. Soc. Middle East 17:13-
17.
Kirchberger, K. 1989. WWF:
WeiBstorchs-Schutzprojekt Rust. Vogelschutz
Oesterreich 4:33-37.
Kirk, J. 1864. On the birds of the Zambesi River
of Eastern Tropical Africa. Ibis VI, 1864:307-
339.
Kirkpatrick, C.E. 1986. Isolation of Salmonella
spp. from a colony of wading birds. J. Wildl.
Dis. 22:262-264.
Kissling, R.E., R.W. Chamberlain, D.B. Nelson
and D.D. Stamm. 1955. Studies on the North
American arthropod-borne Encephalitides.
VII. Equine encephalitis studies in Louisiana.
Am.J. //?£. 62:233-254.
Kist, J., and K.H. Voous. 1962. Avifauna van
Nederland. EJ. Brill., Leiden.
Klaas, E.E., H.M. Ohlendorf, and R.G. Health.
1974. Avian eggshell thickness: variability and
sampling. Wilson Bull. 86:156-164.
Klemm, W. 1989. Status und Schutz des Weiss-
Storchs in Rumaenien. Pp. 125-126 in
Rheinwald,. G., J. Ogden, and H. Schulz (eds),
Weifstorch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Klimov, S.M. 1987. [On nesting of the White
Stork (Ciconia ciconia) in Lipetsk Region.]
Ornitologiya 22:183.
Kling, A. 1963. Zwei neue Vorkommen des
braunen Sichlers (Plegadisfalcinellus) in
Schwaben. Schwaebische Naturhunde (supersedes
Nat. -schaft. Ver. Schwaben.) Berichte 67:61-62.
Klingner, H. 1969. Observing the Glossy Ibis
near Weimar. Falke. Monatsschr. Orn. Vivarienh
Ausg. A. 16:28.
Klos, H.-G. 1970. Berlin Zoo news. Avicult. Mag.
76:204.
Klos, H.-G. 1985. Bemerkungen zur Haltung des
Schuhschnabels im Berliner Zoo. Gejwderte Welt
109:36-37.
Klug, S., and J. Boswall. 1970. Observations
from a water bird colony, Lake Tana,
Ethiopia. Bull. Br. Orn. Club. 90:97-105.
Knierriem, R. 1976. Der Schwarzstorch (Ciconia
nigra) im Vogelsberg. Luscinia 43(l/2):32-33.
Knighton, G.E. 1970. Glossy Ibis in Richmond
County, Georgia. Oriole 35:3.
Knoder, C.E., P.D. Plaza, and A. Sprunt IV.
1980. Status and distribution of the Jabiru
Stork and other water birds in western
Mexico. Pp. 58-127 in Schaeffer, P.P., and
S.M. Ehlers (eds). The Birds of Mexico: Their
Ecology and Conservation. Proceedings of the
353
National Audubon Society Symposium,
Tiburon, California.
Knowlton, F.H. 1909. Birds of the World. Henry
Holt, New York.
Kobayashi, K. 1956a. Birds of Japan in Natural
Colors. Hoikusha, Osaka, Japan.
Kobayashi, K. 1956b. Birds of China. Hoikusha,
Osaka, Japan.
Koch, A., D. Magnus, H. Seilkopf, H. Baron,
and E. Schuz. 1966. Der WeiBstorch-Zug im
Raum Sinai bis Kena in
landschaftsmorphologischer Sicht. Vogelwarte
23:209-220.
Koenig, A. 1910. Die Ergebnisse meiner Reise
nach dem Sudan im Frujahr 1910. Int. Orn.
Kngr. 5:469-545.
Koenig, O. 1952. Okologie und Verhalten der
Vogel des Neusiedlersee—Schilfgurtels. J. Orn.
93:207-289.
Koepcke, H.-W. 1963. Probleme des Vogelzuges
in Perus. Proc. Int. Orn. Congr. 13:396-411.
Koepcke, H.W., and M. Koepcke. 1965. Las aves
silvestres de importancia economica del Peru. Serv.
Forestal y de Gaze, Min. Agric., Lima, Nos. 6
and 7.
Koepke, M. 1970. The Birds of the Department of
Lima Peru. Livingston, Wynnewood,
Pennsylvania, USA.
Kokshaisky, N.V. 1959. [Certain differences
between the Spoonbill (Platalea leucorodia L.)
and the Glossy Ibis (Plegadisfalcinellus L.)
connected with flight.] Dokl. Akad. Nauk SSSR
124:949-952.
Kolbe, H. 1988. Spaetsommerliche Notitzen zur
nordkoreanischen Vogelfauna. Mitt. Zool. Mus.
Berl. 64 suppl.: Ann. Orn. 12:51-66.
Komine, T., and H. Sugita. 1990. Breeding the
Eastern White Stork. Anim. Zoos June:212-
217.
Konig, C. 1982. Zur systematischen Stellung der
Neuweltgeier (Cathartidae). J. Orn. 123:259-
267.
Koningsberger, J.C. 1915. Java, zoologisch en
biologisch. Drukkerij Dep v. L. N. en H.,
Buitenzorg.
Koopman, F.H. \989.Jaarverslag 1989. Aan de
begunstigers van het Ooievaarsbuitenstation
'De Lokkerij' te de Wijk. Spp.
Koppers, N., H. Gerlach, and G. Behlert. 1991.
Mykobakteriose bei Waldrappen (Geronticus
eremita). Berl. Muench. Tieraerztl. Wochenschr.
104:57-62.
Kormos, S. 1929. Ciconia alba mit weissen Lauf.
Aquila 34-35:447-448.
Korsten, J., and H. Lukken. 1990. Gedragsstudie
rode ibis. Stageverslag Hogeschool Gelderland.
Kortegaard, L. 1973. Skestorken (Platalea
leucorodia) i Danmark 1900-1971. [The
Spoonbill (Platalea leucorodia) in Denmark
1900-1971.] Dan. Orn. Foren. Tidsskr. 67(1-
2):3-14.
Kostin, V.Y. 1969. Nesting of (Plegadisfalcinellus
L.) in the Crimea. Vestn. Zool. 1969(1):82-S4.
Kotter, B.L. 1970. An ecological natural history
of the White-faced Ibis (Plegadis chihi) in
northern Utah. M.S. Thesis, Univ. Utah, Salt
Lake City.
Kovacs, B. 1968. Sterile storks on the hortobagy
and in Biharugra. Aquila 75:294.
Kowarski, T., M.A. Pasquier, G. Piette & M.S.
Nouvel. 1937. Recherche d'un parasite
endoglobulaire du genre Haemoproteus chez
diverses especes d'oiseaux du Pare Zoologique
des Bois de Vincennes. Ann. Parasitol. Hum.
Comp. 15:529-536.
Kozlova, E.V. 1932. The birds of South-west
Transbaikalia, Northern Mongolia, and
Central Gobi. Ibis (ser. 13) 2:316-347, 405-
438, 567-596.
Kozlova, E.V., and AJ. Tugarinov. 1947. Fauna
S.S.S.R., nov. ser. no. 33. Ptitsy [Birds], vol. 1,
no. 3. Moscow.
Krabbe, N. 1983. Bald Ibis (Geronticus eremitd)
seen on migration in Israel. OSME Bull. 10:13.
Krai, B. 1965. Functional adaptations of
Ciconiiformes to the darting stroke. Adas Soc.
Zool. Bohemoslov. 29:377-391.
Kramer, H. 1980. Bruten europaischer
Vogelarten in Gefangenschaft. Zool. Garten
(NF) 50:89-137.
Krapivni, A.P. 1958. [On the intraspecific
relationships in White and Black Storks.]
Vestn. Akad. Nauk Belarus. SSR Biol. 1:62-70.
Kreba, R. 1981. Possible sighting of Glossy Ibis
in Saskatchewan. Blue Jay 39:220-222.
Krebs, J.R. 1978. Colonial nesting birds, with
special reference to Ciconiiformes. Pp. 299-314
in Sprunt, A., J.C. Ogden, and S. Winckler
(eds), Wading Birds. Research Report 7,
National Audubon Society, New York.
Krishnan, M. 1960. The Vedanthangal Sanctuary for
Water Birds. Madras State Forest Department,
Madras. 25pp.
Krishnan, M. 1986. The White Stork in Indian
literature. Hornbill 2:24-25.
Kruger, H.-P. 1986. White Stork production
(Ciconia ciconia) from 1979-1984 in the Cottbus
District, East Germany. Beitr. Vogelkde 32:27-
32.
Kuehler, C., and W. Toone. 1989a. The Shoebill:
shy creature of the swamps. Zoonooz May.
Kuehler, C., and W. Toone. 1989b. Zoological
Society of San Diego Shoebill Stork
management plan (Balaeniceps rex}.
Unpublished report, San Diego Zoological
Society, San Diego, California.
Kuerzi, J.F. 1931. Notes on the occurrence of
Ajaia ajaja and some other species in Florida.
Auk 48:114-116.
Kuerzi, R.M. 1938. Some notes on the birds of
St. Lucie Co., Florida. Fla. Nat. 12:11-15.
Kuhk, R. 1954. Legenden und Falschmeldungen
ueber Storch. Vogelwarte 17:165-166.
Kuhk, R. 1956. Hagelunwetter als
Verlustursache bei Storche und anderen
Vogeln. Vogelwarte 18:180-182.
Kuhk, R. 1960. Brutvorkommen von WeiBstorch
(C. ciconia) und Braunem Sichler (Plegadis
falcinellus] in Oberitalien. Vogelwarte 20:292-
293.
Kuhk, R., and K. Hirschfeld. 1957. Zwei 17
jahrige WeiBstorche (C. ciconia). Vogelwarte
19:143-144.
Kuhk, R., and R. Schuz. 1950. 1949
Storungsjahr im Bestand des WeiBstorche
Ciconia ciconia. Orn. Beob. 47:93-97.
Kuhk, R., and R. Schuz. 1956. Ein Nortruf:
Weisser Storch in Gefahr! Pp. 165-169 in
Steiniger, Natur undjagd in Niedersachsen
(Festschrift Hugo Weigold). Hannover.
Kumar, C.S. 1982. Ecology and behaviour of the
Indian Black Ibis, Pseudibispapillosa
(Temminck). M.S. Dissertation, University of
Saurastra, India.
Kumar, C.S., and V.C. Soni. 1984. Laboratory
observations on the incubation period of the
Indian Black Ibis (P. papillosa Temmink).y.
Bombay Nat. Hist. Soc. 81:189-191.
Kumerloeve, H. 1958. Von der Kolonie des
Waldrapps, Geronticus eremita (L.), bei Birecik
am Euphrat. Beitr. Vogelkd. 6:189-202.
Kumerloeve, H. 1960. On some birds collected
by Mr. Douglas Carruthers in the Syrian
Desert. Alauda 28:284-286.
Kumerloeve, H. 1961. LaCigogne blanche,
Ciconia c. ciconia (L.), en Anatolic, Syrie, Liban
et Palestine. Alauda 29:25-30.
Kumerloeve, H. 1962. Zur Geschichte der
Waldrapp-Kolonie in Birecik am oberen
Euphrat. J. Orn. 103:389-398.
Kumerloeve, H. 1964a. Ungewohnliche
Verhaltensweisen bei Reihern und Storchen.
Orn. Mitt. 16:59.
Kumerloeve, H. 1964b. ZurSumpf- und
Wasservogelfauna der Turkei. J. Orn.
105:307-325.
Kumerloeve, H. 1965. Zur Situation der
Waldrappkolonie (Geronticus eremitd) (L. 1758)
in Birecik am Euphrat. Vogelwelt 86:42-48.
Kumerloeve, H. 1966a. Zur Brutverbreitung und
Durchzug des WeiBstorches C. ciconia (L.) in
Kleinasien. Vogelwarte 23:221-224.
Kumerloeve, H. 1966b. Zum Zug des
Schwarzstorches Ciconia nigra (L.) in der
Turkei. Vogelwarte 23:310-311.
Kumerloeve, H. 1967. Nouvelles donnees sur la
situation de la colonie d'Ibis chevelus
Geronticus eremita (L.) 1758 a Birecik sur
L'Euphrate (Turquie). Alauda 35:194-202.
Kumerloeve, H. 1969a. Vom Waldrapp,
Geronticus eremita (L.,1758), dem einstigen
Brutvogel der Alpen.Jahrb. Ver. Schutze
Alpenpfanz. Tiere 34:132-138.
Kumerloeve, H. 1969b. Situation de la colonie
d'Ibis chevelus Geronticus eremita a Birecik en
1968 et 1969. Alauda 37:260-261.
Kumerloeve, H. 1970. On the composition and
direction of fall migration in 1969 on the
Montenegrin Littoral. Larus 24:119-124.
Kumerloeve, H. 1971. Zum Brutvorkommen des
Sichlers (Plegadis falcinellus L.) im vorderen
Orient. Zool. Abh. (Dres.) 30(19):243-246.
Kumerloeve, H. 1972. Vom Waldrapp, Geronticus
eremita (L, 1758) —Einige Bemerkungen zu
seiner Entdeckungsgeschichte.yaAr^. Ver.
Schutze Alpenpflanz- Tiere 37:1-6.
Kumerloeve, H. 1978a. Bemerkungen zur
Situation von Waldrapp und Kahlkopfibis.
Angew. Orn. 4:114-116.
Kumerloeve, H. 1978b. Waldrapp, Geronticus
eremita (Linnaeus, 1758), und Glattnackenrapp,
Geronticus calvus (Boddaert,1783): Zur
Geschichte ihrer Erforschung und zur
gegenwartigen Bestandssituation. Ann.
Naturhist. Mus. Wien 81:319-349.
Kumerloeve, H. 1983. Zur Kenntnis
altagyptischer Ibis-Darstellungen, unter
besonderer Beriicksichtigung des Waldrapps,
Geronticus eremita (Linnaeus, 1758). Bonn., Zool.
Beitr. 34:197-234.
Kumerloeve, H. 1984. The Waldrapp, Geronticus
eremita (Linnaeus, 1758): historical review,
taxonomic history, and present status. Biol.
Conserv. 30:363-373.
Kumerloeve, H. 1988. Zur Brutsituation des
Weiss-storchs, Ciconia ciconia, in Anatolien
(Stand 1987). [The breeding situation of
White Stork Ciconia ciconia, in Anatolia (Stand
1987).] Bonn Zool. Beitr. 39:361-370.
Kumerloeve, H. 1989. Status und Verbreitung
des WeiBstorchs in der Turkei 1958/1987. Pp.
175-176 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weifstorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Kurochkin, E.N. 1982. [New birds from the
Middle Pliocene of Mongolia.] Ornitologiya
17:150-154.
Kuroda, N. 1936. Birds of the Island of Java.
Tokyo.
Kuroda, N. 1970. Ende der japanischen
Population des Schwarzschnabel-Storchs
(Ciconia ciconia boyciana). Vogelwarte 25:269.
Kushlan, J.A. 1973a. White Ibis nesting in the
Florida Everglades. Wilson Bull. 85:230-231.
Kushlan, J.A. 1973b. Promiscuous mating
behavior in the White Ibis. Wilson Bull.
85:331-332.
Kushlan, J.A. 1976a. Feeding behavior of North
American herons. Auk 93:86-94.
Kushlan, J.A. 1976b. Wading bird predation in a
seasonally fluctuating pond. Auk 93:464-476.
Kushlan, J.A. 1976c. Feeding rhythm in nestling
White Ibis. Wilson Bull. 88:656-658.
Kushlan, J.A. 1976d. Site selection for nesting
354
colonies by the American White Ibis Eudocimus
albus in Florida. Ibis 118:590-593.
Kushlan, J.A. 1977a. Foraging behavior of the
White Ibis. Wilson Bull. 89:342-345.
Kushlan, J.A. 1977b. Growth energetics of the
White Ibis. Condor 79:31-36.
Kushlan, J.A. 1977c. Population energetics of the
American White Ibis. Auk 94:114-122.
Kushlan, J.A. 1977d. Sexual dimorphism in the
White Ibis. Wilson Bull. 89:92-98.
Kushlan, J.A. 1977e. The significance of
plumage colour in the formation of feeding
aggregations of Ciconiiformes. Ibis 119:361-
364.
Kushlan, J.A. 1977f. Differential growth of body
parts in the White Ibis. Auk 94:164-166.
Kushlan, J.A. 1978a. Feeding ecology of wading
birds. Pp. 249-297 in Sprunt, A., J.C. Ogden,
and S. Winckler (eds), Wading Birds. Research
Report 7, National Audubon Society, New
York.
Kushlan, J.A. 1978b. Commensalism in the
Little Blue Heron. Auk 95:677-681.
Kushlan, J.A. 1978c. Nonrigorous foraging by
robbing egrets. Ecology 59:649-653.
Kushlan, J.A. 1978d. Wading bird use of the east
Everglades. Fla. Field Nat. 6:46-47.
Kushlan, J.A. 1979a. Design and management of
continental wildlife reserves: lessons from the
Everglades. Biol. Conserv. 15:281-290.
Kushlan, J.A. 1979b. Prey choice by tactile-
foraging wading birds. Proc. Colonial Waterbird
Group 3:133-142.
Kushlan, J.A. 1979c. Feeding ecology and prey
selection in the White Ibis. Condor 81:376-389.
Kushlan, J.A. 1979d. Wading bird preservation
in Everglades National Park. Pp. 483-487 in
Proceedings of the First Conference on Scientific
Research in National Parks. US Department of
Interior, Washington, D.C.
Kushlan, J.A. 1979e. Effects of helicopter
censuses on wading bird colonies. J. Wildl.
Manag. 43:756-760.
Kushlan, J.A. 1979f. White Ibis. Pp. 83-84 in
Rare and Endangered Biota of Florida. Vol. 2. Birds.
University Presses Florida, Gainesville,
Florida.
Kushlan, J.A. 1980. Reintroduction of
indigenous species to natural ecosystems.
Environ. Manag. 4:93—94.
Kushlan, J.A. 1981. Resource use strategies of
wading birds. Wilson Bull. 93:145-163.
Kushlan, J.A. 1983. Special species and
ecosystems preserves: colonial waterbirds in
the U.S. national parks. Environ. Manag. 7:201-
207.
Kushlan, J.A. 1985a. Storks and spoonbills. Pp.
72-83 in Perrings, C.M., and A.L.A.
Middleton (eds), Encyclopedia of Birds. Facts-
on-File Publishers, New York, and Unwin,
London.
Kushlan, J.A. 1985b. Review: recovery plan for
the U.S. breeding population of the Wood
Stork. Colonial Waterbirds 10:259-262.
Kushlan, J.A. 1986a. Colonies, surveys, and
sites: the terminology of colonial waterbird
research. Colonial Waterbirds 9:119-120.
Kushlan, J.A. 1986b. Responses of wading birds
to seasonally fluctuating water levels:
strategies and their limits. Colonial Waterbirds
9:155-162.
Kushlan, J.A. 1987. External threats and
internal management: the hydrologic
regulation of the Everglades. Environ. Manag.
11:109-119.
Kushlan, J.A. 1989a. Avian use of fluctuating
wetlands. Pp. 593-604 in Sharitz, R.R., and
J.W. Gibbons (eds), Freshwater Wetlands and
Wildlife. DOE Symposium Series No. 61. US
DOE, Oak Ridge, Tenn.
Kushlan, J.A. 1989b. Wetlands and wildlife, the
Everglades perspective. Pp. 773-790 in
 Sharitz, R.R., and J.W. Gibbons (eds),
Freshwater Wetlands and Wildlife. DOE
Symposium Series No. 61. US DOE, Oak
Ridge, Tenn.
Kushlan,J.A. 1990a. Freshwater marshes. In
Meyers, R.L., and J.J. Ewel (eds), Ecosystems of
Florida. University of Central Florida Press,
Orlando.
Kushlan, J.A. 1990b. The Everglades system. In
Livingston, RJ. (ed.), Streams of Florida.
Springer-Verlag, New York.
Kushlan, J.A., and O.L. Bass, Jr. 1983.
Decreases in the southern Florida Osprey
population, a possible result of food stress.
Pg.181-200 in Bird, D.M. (ed.), Status and
Management of Osprey and Eagles. Raptor
Research Foundation, Montreal.
Kushlan, J.A., and K.L. Bildstein. in press.
White Ibis. In Poole, A., P. Stettenheim, and
F. Gill (eds.) Birds of North America. American
Ornithologist's Union, Philadelphia,
Pennsylvannia.
Kushlan, J.A., and P.C. Frohring. 1985.
Decreases in the Brown Pelican population in
Southern Florida. Colonial Waterbirds 8:83-95.
Kushlan, J.A., and. P.C. Frohring. 1986. History
of the southern Florida Wood Stork
population. Wilson Bull. 98:368-386.
Kushlan, J.A., and M.S. Kushlan. 1975. Food of
the White Ibis in southern Florida. Fla. Field
Nat. 3:31-38.
Kushlan, J.A., and W.B. Robertson, Jr. 1977.
White Ibis nesting in the lower Florida Keys.
Fla. Field Nat. 5A\^2.
Kushlan, J.A. andJ.L. Schortmeyer. 1974.
Glossy Ibis nesting in southern Florida. Fla.
Field Nat. 2:13-14.
Kushlan, J.A., and D.A. White. 1977. Nesting
wading bird populations in southern Florida.
Fla. Set. 40:65-72.
Kushlan, J.A., J.C. Ogden, and A.L. Higer.
1975. Relation of water level and fish availability to
Wood Stork reproduction in the southern Everglades,
Florida. U.S. Geological Survey, Open File
Report. 75-434. Tallahassee, Fla.
Kushlan, J.A., M.C. Baumann, and L.C.
McEwan 1979. A bibliography of southern Florida
wading birds. South Florida Research Center
Report, National Park Service, Homestead,
Florida.
Kushlan, J.A., L.G. Morales and P.C. Frohring.
1985. Foraging niche relations of wading birds
in tropical wet savannahs. Pp. 663-682 in
Buckley, P.A., M.S. Foster, E.S. Morton, R.S.
Ridgley, and F.G. Buckley (eds), Neotropical
Ornithology. Ornithological Monograph No. 36.
American Ornithologists' Union, Washington
D.C.
Kwak, J.-Y., T.H. Koo, and P.-O. Won. 1980.
Breeding ecology of the White Stork, Ciconia
boyciana Swinhoe in Korea. Bull. Korean Assoc.
Conserv. Nat. ser. 11:105-117.
Kyle, J.L. 1933. White Ibises at Orange Lake,
Florida. Fla. Nat. 6:45-47.
Kyllingstad, H.C. 1986. A record of Bald Ibis
from the Sinai Mountains. Bull. Orn. Soc.
Middle East 17:1-2.
LaBastille, A. 1974. Taj Mahal to nature.
Audubon 76:20-33.
Lack, D. 1954. The Natural Regulation of Animal
Numbers. Clarendon Press, Oxford. 343pp.
Lack, D. 1966. Population Studies of Birds.
Clarendon Press, Oxford. 341pp.
Lack, D. 1968a. Ecological Adaptations for Breeding
in Birds. Methuen, London.
Lack, D. 1968b. Bird migration and natural
selection. Oikos 19:1-9.
Lack, D., and A. Lack. 1973. Birds on Grenada.
7£w 115:53-59.
Laedlein-Greilsammer, D., J.M. Lernould, and
C. Peters. 1979. Tumeurs hepatiques chez les
cigognes blanches (Ciconia ciconia) et les
Cigognes noires (Ciconia nigra) au Pare
Zoologique et Botanique de Mulhouse. Pp.
239-242 in Verhandlungsbericht des XXL
International Symp. tiber die Erkrankungen der
Zootiere Mulhouse, 1979.
Lalas, C. 1974. Australian Ibis. Wildl. Aust.
11:30-32.
Lamb, C. 1922. Some birds recently observed in
southern California. Condor 24:184.
Lambert, F. 1983. Report on a brief
ornithological visit to Trinidad. Unpublished
report, International Council for Bird
Preservation, Cambridge, England.
Lambert, F.R., and R.F. Grimmett. 1983. Bald
Ibis (Geronticus eremita] in Israel. OSME Bull.
10:12.
Land, H.C. 1970. Birds of Guatemala. Livingstone
Publishing Company, Wynnewood,
Pennsylvania.
Lange, H. 1940. Storkens Alder og Yngleforhold i
Denmark. Munksgard, Kobenhavn.
Lange, H. 1942. Storkens udbredelse.
Storkebygder i Danmark og tal-svinghinger i
Europa. Dan. Orn. Foren. Tidsskr. 36:70-92.
Lange, H. 1948. Storken (Ciconia ciconia (L.)) i
Sydvest-Jylland 1940-1947. Dan. Orn. Foren.
Tidsskr 42:4-10.
Lange, H. 1954. Gibt es zweijahrige Bruter und
alte Nichtbruter beim Weissen Storch?
Vogelwarte 17:150-155.
Lange, H. 1974. Novemberbeobachtung eines
WeiBstorches. Falke 21:176.
Lange, H., W. Emeis, E. Schuz, and R. Drost.
1938. Weitere Angaben uber Heimkehr-
Verzogerung und Bestand des Weissen
Storches 1937. Vogelzug 9:97-102.
Langrand, O. 1990. Guide to the Birds of
Madagascar. Yale University Press, New
Haven.
Lantz, D.E. 1899. A list of birds collected by Col.
N.S. Goss in Mexico and Central America.
Trans. Kans. Acad. Sci. 16:218-224.
Lantz, D.E. 1900. The Roseate Spoonbill in
Kansas. Auk 17:171.
Lantz, D.E. 1907. An economic study ofjield mice.
USDASurv. Bull. 31.
Lassus, N.S. 1973. Saddle-bill Storks. Bull. E.
Afr. Nat. Hist. Soc. (May, 1973):74.
Latham, J. 1790. Index ornithologicus, sive Systema
Ornithologiae, vol. 2. Harvard University
Museum of Comparative Zoology.
Latham, J. 1809. Systema Ornithologiae sive Index
Ornithologicus. Johanneau, Paris.
Lathbury, G., 1970. A review of the birds of
Gibraltar and its surrounding waters. Ibis
112:25-43.
Lathigara, R. 1989. Behavioural and ecological
studies on population census, roosting,
foraging, feeding, and breeding of the Indian
Black Ibis at Jamnagar. M.S. Dissertation,
Saurashtra University, Rajkot, India.
La Touche, J.D.D. 1920. Notes on the birds of
north-east Chihli in north China. Ibis
ll(2):629-671,880-920.
La Touche, J.D.D. 1921. Notes on the birds of
north-east Chihli, in North China. Part III.
Ibis (ser. 11)3:3-48.
La Touche, J.D.D. 1931-1934. A Handbook of the
Birds of China, revised 5th edn, volume 2.
Taylor and Francis, London.
Laubmann, A. 1939. Die Voegel von Paraguay, vol.
1. Strecker and Schroeder, Stuttgart.
Lauhachinda, V. 1969. Preliminary study of the
life history of the Open-billed Stork, Anastomas
oscitans (Boddaert). M.S. Thesis, Kasetsart
University, Bangkok, Thailand.
Laukel, H. 1987. Ein Traum in Rot. Das Tier
4:10-13.
Lauthe, P. 1976. Vom Bestand des WeiBstorches
(Ciconia c. ciconia) in Tunesien bis 1975.
Gefoderte Welt WQ:\38.
355
Lauthe, P. 1977. La cigogne blanche en Tunisie.
Oiseau Rev. Fr. Orn. 47(3): 223-242.
Lauthe, P. 1989. Status, Verbreitung und Zug
des WeiBstorchs in Tunesien. Pp. 189-194 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weif storch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Lavery, HJ. 1971. Wild Ducks and other Waterfowl
in Queensland. Department Primary Industries,
Brisbane. 58pp.
Lavery, H.J., D. Seton, and J.A. Bravery. 1968.
Breeding seasons of birds in north-eastern
Australia. Emu 68:133-147.
Law, J.E. 1912. Wood Ibis near Long Beach.
Condor 14:41.
Law, S.C. 1926. The nesting of the Open-bill
Stork (Anastomus oscitans) in Purulia,
Manbhum District. J. Bombay Nat. Hist. Soc.
31:223-226.
Lawrence, J.W. 1979. White Ibis in Wilson
County. Migrant 50:61.
Lawson, W J. 1965. On the record of the White
Stork Ciconia ciconia (Linnaeus) breeding in
Zululand. Ostrich 36:140.
Lawton, J. 1989. Last of the Mohicans. Aramco
world 4(6):2-5.
Layard, E.L. 1867. The Birds of South Africa. Juta,
Cape Town.
Layard, E.L., and R.B. Sharpe. 1884. The Birds of
South Africa. Quaritch, London.
Laycock, H.T. 1965. Aquatic birds at Blantyre.
(tar^ 36: 123-1 30.
Lazaro, E. 1986. Beitrag zur Ernahrungsbiologie
des WeiBstorchs in Spanien. Beth. Veroff
Naturschutz Landschaftsplege Bad.-Wurtt 43:235-
242.
Lazaro, E., and M. Fernandez-Cruz. 1989.
Migration of the White Stork through the
Strait of Gibraltar in 1985: first results. Pp.
263-268 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weifistorch: Status und Schutz, Proc.
1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Lazaro, E., P. Chozas, and M. Fernandez-Cruz.
1986. [Breeding population of the White Stork,
C. ciconia, in Spain. National census 1984.]
Ardeoia 33:131-169.
Leach, J. A., and P.C. Morrison. 1958. An
Australian Bird Book, 9th edn. Whitcombe and
Tombs, Melbourne. 224pp.
Lebedeva, M.I. 1958. [The White Stork in
Azerbaijan.] Priroda (Mosc.) 9:104-105.
Lebedeva, M.I. 1959a. [Biology of the Black
Stork (Ciconia nigra) in the Belovezskoi forest.]
Ornitologiya 2:138-142.
Lebedeva, M.I. 1959b. [Register of the number
of White Storks in the Soviet Union.] All-Soviet
Orn. Conf. 2:26-28.
Lebedeva, M.I. 1960a. [On the number of White
Storks in the Soviet Union.] Ornitologiya 3:413-
419.
Lebedeva, M.I. 1960b. [Some information on the
migration and numbers of the White Stork.]
Migracii Schivotnych 2, Akad. Nauk SSSR 2:130-
139.
Lebedeva, M.I. 1976. Numbers, migration, and
ecology of storks in the USSR, Pp. 1-25.
(Synopsis of dissertations prepared for
candidate of Biological Sciences Degree.)
University of Moscow.
Lebedeva, M.I. 1977a. [The range and numbers
of Oriental White Storks in the USSR.] Pp.
228-229 in Winter, S. (ed.) VII All-Soviet
Ornithological Conference. Zoological Institute of
the Academy of Science of the Ukranian SSR,
Kiev.
Lebedeva, M.I. 1977b. Distribution and
population of the Far Eastern Stork in USSR.
USSR Nat. Orn. Conf. Kiev 7:299.
Lebedeva, MJ. 1989. Verbreitung und
 Bestanddynamik des WeiBstorchs in der
russischen S.F.S.R. Pp. 159-160 in Rheinwald,
G., J. Ogden, and H. Schulz (eds), Weiss-
Storch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Lebedeva, M.I., and T.P. Schevareva. 1960.
[Causes of death of birds in nature (on the
basis of banding results).] Bull. Ochrana Prirody
i Sapowednoe Delo SSSR 6:67-83.
Leber, K.K. 1980. Habitat utilization in a
tropical heronry. Brenesia 17:97-136.
Ledant,J.P.,J.P.Jacob, P.Jacobs, F. Mahler, B.
Ochando, and J. Roche. 1981. Mise a jour de
1'avifaune algerienne. Gerfaut 71:295-398.
Lee, D.S. 1987. Breeding birds of Carolina bays:
succession-related density and diversification
on ecological islands. Chat 51:85-102.
Leenhouts, W.P. 1986. Wood Storks feeding on
fish from dredging activities. Fla. Field. Nat.
15:22.
Lefranc, N., and B. Schmitt. 1969. Donnees
inedites sur 1'avifaune alsacienne du Nord de
Strasbourg et sur son evolution de 1958 a
1968. Alauda 37:224-229.
Legany, A. 1979. Data on the bird stock of the
floodplain at Tiszafured, Hungary. Tiscia
(Szeged) 13:157-162.
Legge, W.V. 1880. A History of the Birds of Ceylon.
Tisaria, Dehiwala, Sri Lanka.
Legler, K. 1987. Plegadis ibis near Goose Pond in
Columbia County. Passenger Pigeon 49:160.
Leja, P. 1989. [Our friend-the White Stork.]
PutniDaba 2:189-190.
Lekagul, B. 1968. Bird Guide of Thailand. Assoc.
Conserv. Wildl., Bangkok. 272pp.
Lekagul, B., and E.W. Cronin, Jr. 1974. Bird
Guide of Thailand, 2nd edn. Assoc. Conserv.
Wildl., Bangkok. 324pp.
Leonovich, V.V., and L.A. Nikolyevski. 1976.
[Notes about Far Eastern Stork breeding.]
Trans. Oka State Nat. Res. 12:17-19.
Leotaud, A. 1866. Oiseaux de I'He de la Trinidad
(Antilles). Chronicle publishing office, Port
d'Espagne.
Lepasaar, J. 1985. White storks produce a repeat
clutch. Eesti Loodus 28:514, 541, 543.
Leshem, Y. 1990. [On a White Stork Ciconia
ciconia from Germany caught in Israel in 1846.]
Torgos 8:55, 76.
Lesniczak, A.B. 1989. The Black Stork, Ciconia
nigra, in the environs of Rybnik on the Silesian
Upland. Chronmy Przyr. Ojczysta 45:68-70.
Le Souef, W.H.D. 1917. Australian ibises. Emu
17:94-95.
Lesser, F.H. 1964. Waterbirds of Little Creek.
Cassinia 6:22-32.
Leuthold, W., and B. Leuthold. 1972.
Blutschnabelweber (Quelea quelea) als Beute
van Greif- und Stelzvoegeln. Vogelwarte
26:352-354.
Leveque, R. 1960. Observations de Espatulas en
Huelva, Abril-Mayo 1959. Ardeola 6:378-379.
Leveque, R. 1964. Notes on Ecuadorian birds.
Ibis 106:52-62.
Leveque, R. 1966. Sobre avifauna de Portugal en
invierno. Ardeola 11:101-107.
Lewis, A.D., and DJ. Pearson. 1981a. Raptor
and stork migration at Namanga, southern
Kenya. Scopus 5:83-84.
Lewis, A.D., and DJ. Pearson. 1981b. Stork and
raptor migration in South Nyanza, Kenya.
Scopus 5:51.
Lewis, C.B. 1948. Unverified observations of
'dozens of Glossy Ibis and a number of White
Ibis ... feeding at the mouth of the Duhaney
River. Nat. Hist. Notes Nat. Hist. Soc. Jamaica
32:142.
Lewis, J.C. 1983. Habitat suitability index models:
Roseate Spoonbill. U.S. Fish Wildl. Serv. Biol.
Rep. FWS/OBS-82:10.50. 16pp. US Dept of
the Interior, Washington DC.
Lewis, T.H. 1971. Field notes on the dry season
birds of Nayarit. Tex.J. Set. 23:57-66.
Lewis, T.H. 1972. Notes on the dry season birds
ofSinaloa. Tex.J. Sci. 24:233-243.
Li, W.H., H. Zhao, and X. Luan 1989.
[Reproductive ecology of Oriental Stork and
feeding of young birds.]J. Heilongjiang August
First Land Reclam. Univ. 2:9-18.
Li, W. et al. 1990. [Reproductive ecology and
taming of Oriental White Storks.] Chinese
Wildl. 1990(2): 16-17, 27.
Li, W., H. Zhao, and X. Luan. 1991.
Reproductive ecology of the Oriental White
Stork (Ciconia boyciand) with information on
feeding and development of the chicks. Pp. 47-
58 in Coulter, M.C., Q. Wang, and C.S.
Luthin (eds), Biology and Conservation of the
Oriental White Stork (Ciconia boyciana).
Savannah River Ecology Laboratory, Aiken,
South Carolina, USA.
Libbert, W. 1954. Wo verbleiben die WeiB-
storche aller Altersstufen in den Brutmonaten?
Voglewarte 17:100-113.
Libbert, W. 1971. Ein Schwarzstorchpaar siedelt
sich in einem uckermarkischen Waldrevier an.
Falke 18:58-60.
Libbert, W., H. Ringleben, and E. Schuz. 1937.
Ring-Wiederfunde deutscher Weisz-Storch (C.
c. ciconia) aus Afrika und Asien. Vogelzug 8:193-
208.
Lieu Yin-zeng. 1981. [Discovery of Japanese
Crested Ibis in Qin-Ling.] Acta Zoo/. Sinica
27:273.
Ligon, J.D. 1967. Relationships of the cathartid
vultures. Occas. Pap. Mus. Zoo/. Univ. Mich.
651:1-26.
Limpricht, J.A. 1717. Ciconiae anatome.
Leopoldina 5-6:209-212.
Lincoln, F.C. 1923. The White Ibis in California.
Co«</or25:181.
Lindblad, J. \969.Journey to Red Birds. Hill and
Wang, New York.
Linder, R. 1977. Die Entwicklung der Weiss-
Storchpopulation (Ciconia ciconia) im Landkreis
Dieburg von 1945 bis zum erloschen. Luscinia
43(3/4): 115-123.
Line, LJ. 1941. Nesting of the Hadada Ibis
(Hagedashia hagedash hagedash). Ostrich 11:137-
139.
Line, L. 1971. The storks come back. Audubon
73:26-27.
Linnaeus, C. 1758. Systema Naturae, 10th edn.
Salvius, Holmiae.
Lippens, L. 1938. Les Oiseaux aquatiques du
Kivu. Gerfaut (Fasc. Spec.) 28:1-119.
Lippens, L. 1954. Les Oiseaux d'Eau de Belgique. E.
Vercruysse, Bruges. 302pp.
Lippens, L., and H. Willie. 1972. Atlas des Oiseaux
de Belgique et d'Europe occidentale. Tielt.
Lippens, L., and H. Willie. 1976. Les Oiseaux du
Zaire. Edit. Lannoo Tielt, Belgium. 509pp.
Lippert, W. 1970. Der Indien-Nimmersatt. Falke
17:214.
Lisle, M. d. 1956. Sur une observation de
Balaeniceps rex au Cameroun. Oiseau Rev. Fr.
Orn. 26:1-3.
Little, VJ. d. 1967a. Some aspects of the
behaviour of the Wattled Plover (L.). Ostrich
38:254-259.
Little, VJ. d. 1967b. Sight record of an Openbill.
Ostrich 38:204.
Litvinenko, N.M. 1967. Der
Schwarzschnabelstorch (Ciconia ciconia boyciana)
im Bezirk Chabarowsk. Pp. 188-189 in Schutz,
rationelle Nutzung und Erneuerung der naturlichen
Hilfsquellen des Amurlandes. Thesen der
Wissenschaftl. Konferenz Chabarowsk.
Litvinenko, N.M. 1968. On distribution of the
Oriental White Stork at Lower Amur.
Ornitologiya 9:356-358.
Liversidge, R. 1955. Knob-billed Goose,
356
Spoonbill, and Little Swift from the vicinity of
Cape Town. Ostrich 26:45.
Lloyd, C.A. 1895. Stray notes from Pirara.
Timehri 9:220-232.
Lloyd, C.A. 1897. Nesting of some Guiana birds.
Timehri \ 1:1-10.
Locatelli, M.S. and T.E. Blankenship. 1973.
White-faced Ibis in Humbolt County,
California. Murrelet 54: 1 1 .
Lockhart, P.S. 1975. Glossy Ibis fishing
technique. Bokmakierie 27:84.
Loftin, R.W., and R.D. Bowman. 1978. A device
for measuring egg volumes. Auk 95:190-191.
Lohmer, R. 1989. Kritische Anmerkungen zur
Beringung des WeiBstorchs. Pp. 425-434 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weifstorch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Lohrl, H. 1957. WeiBstorch 'erschnabelt' Beute
im Wasser. Vogelwarte 19(l):52-53.
Lohrl, H. 196 la. Verhaltensweisen eines
erfahrungslosen Weissen Storches. Vogelwarte
21:137-142.
Lohrl, H. 1961b. Ciconia ciconia (Ciconiidae):
Nahrungserwerb. Encyclop. Cinematogr.,
Farbfilm E 375/1961. Inst. Wissensch. Film,
Gottingen.
Lohrl, H. 1968. Das Nesthakchen als
biologisches Problem./ Orn. 109:383-395.
Loiseau, AJ. 1977. Nidification de la cigogne
noire (Ciconia nigra) en Franche-Comte:
premiere donnee Francaise. Alauda 45:335-
338.
Lokemoen, J.T. 1979. The status of herons,
egrets, and ibises in North Dakota. Prairie Nat.
Long, A.J., et al. 1988. A Survey of Coastal Wetlands
and Shorebirds in South Korea, Spring 1988. Asian
Wetland Bureau, Kuala Lumpur. 163pp.
LongJ.L. 1981. Introduced Birds of the World.
David and Charles, Newton Abbot.
Long, R.C. 1967. The birds of the Port Herald
District. Ostrich 38:37-45.
Longstreet, RJ. 1929. Great White Heron and
Roseate Spoonbill near Daytona Beach,
Florida. Auk 46:105.
Longstreet, RJ. 1933. Roseate Spoonbill (Ajaia
ajaja) in Mosquito Lagoon. Fla. Nat. 7:16.
Lonnberg, E. 1903. On a collection of birds from
northwestern Argentina and the Bolivian
Chaco. Ibis (ser. 8) 3:441-471.
Lonnberg, E. 1911. Birds collected by the
Swedish Zoological Expedition to British East
Africa. Kungl. Svenska Vetenskapsakademiens
£//it»///tW/ir 47-1-131
Lopez O.A., J.F. Lynch, and B. MacKinnon de
Monies. 1989. New and noteworthy records of
birds from the eastern Yucatan Peninsula.
Wilson Bull. 101:390-409.
Lorber, P. 1982. Further notes on the Black
Stork in Zimbabwe. Honeyguide 110:8-14.
Louette, M. 1988. Additions and corrections to
the avifauna of Zaire (3). Bull. Br. Orn. Club
108:112-120.
Love, G.J., S.A. Wilkin, and M.H. Goodwin, Jr.
1953. Incidence of blood parasites in birds
collected in southwestern Georgia. J. Parasitol.
39:52-57.
Loveridge, A. 1960. An avian visitor. Ostrich
31:26.
Lowe, D.W., J.R. Matthews, and CJ. Moseley,
(eds.) 1990. Wood Stork, Mycteria americana.
Pp. 653-655 in The Official World Wildlife Fund
Guide to Endangered Species in North America.
World Wildlife Fund/Beacham Publ.,
Washington, DC.
Lowe, K.W. 1982. Feeding behavior and diet of
Royal Spoonbills Platalea regia in Westernport
Bay, Victoria, Australia. Emu 82:163-168.
Lowe, K.W. 1983. Egg size, clutch size, and
 breeding success of the Glossy Ibis Plegadis
falcinellus. Emu 83:31-34.
Lowe, K.W. 1984. The breeding biology of the
Sacred Ibis Threskiornis aethiopicus in Southern
Victoria. PhD Dissertation. Univ. of
Melbourne, Parkville, Australia.
Lowe, K.W. and G.C. Richards. 1991.
Morphological variation in the Sacred Ibis
Threskiornis aethiopicus superspecies complex.
Emu. 91:41-45.
Lowe, K.W., A. Clark, and R.A. Clark. 1985.
Body measurements, plumage and moult of
the Sacred Ibis in South Africa. Ostrich
56:111-116.
Lowe, V.T. 1957. Glossy Ibis notes. Emu 57:347-
348.
Lowe, W.P. 1894. The Scarlet Ibis (Guam rubra)
in Colorado. Auk 11:224.
Lowery, G.H., Jr. 1955. Louisiana Birds.
Louisiana State University Press, Baton
Rouge.
Lowery, G.H., Jr., and W.W. Dalquest. 1951.
Birds from the state of Veracruz, Mexico.
Univ. Kans. Publ. Mus. Nat. Hist. 3:531-649.
Lowther, E.H.N. 1949. A Bird Photographer in
India. Oxford University Press, Oxford,
England.
Loyn, R.H. 1975. Report on the avifauna of
Westernport Bay. Melbourne Min. Conserv.,
Melbourne.
Loyn, R.H. 1978. A survey of birds in
Westernport Bay, Victoria, 1973-74. Emu
78:11-19.
Lucanus, F. v. 1931. Die Zugwege des Weissen
Storchs und des Schwarzstorchs. Proc. Int. Orn.
Congr. 7:327-332.
Lucas, A.H.S., and W.H. D. Le Souef. 1911. The
Birds of Australia. Whitcombe and Tombs,
Melbourne.
Lucker, J.T. 1941. A redescription of Contracaecum
multipapillatum (Von Drasche, 1882)
(Nematoda:Anisakinae). J. Parasitol. 27:505-
512.
Luders, L. 1953. Katastrophenjahr fur Storche.
Heimatfreund 104(30).
Lumsden, R.D. 1961. The White Ibis Eudocimus
albus (Linn.), host for the diplostomatic
trematode Fibricola cratera (Barker & Noll,
1915) Dubois, 1932, in Louisiana./ Parasitol.
17:897.
Lumsden, R.D. 1962. Four echinostome
trematodes from Louisiana birds including the
description of a new species. Tulane Stud. Zool.
9:301-308.
Lumsden, R.D. and J.A. Zischke. 1963. Studies
on the trematodes of Louisiana birds. Zeitschr.
Parasitol. 22:316-366.
Luthin, C.S. 1983a. Observations on the
breeding ecology and behavior of three
Neotropical ibises (Threskiornithidae) on a
ranch in Central Venezuela. M.S. Thesis,
University of Wisconsin, Madison.
Luthin, C. 1983b. Breeding ecology of
Neotropical ibises in Venezuela and comments
on captive propagation. Pp. 95-124 in
Proceedings of the Jean DelacourllFCB Symposium.
International Foundation for the Conservation
of Birds, Hollywood, California.
Luthin, C.S. 1983c. World working group on
storks, ibises, and spoonbills, Report 1,
Summer 1983. Supplement to flying Free 1(1),
Newsletter to the W. W. Brehm-Fundfor
International Bird Conservation, Vogelpark
Walsrode, Walsrode, West Germany.
Luthin, C.S. (ed.) 1984a. World working group on
storks, ibises, and spoonbills, Report 2,1984.
International Council for Bird Preservation,
W.W. Brehm Fund for International Bird
Conservation, Vogelpark Walsrode, Walsrode,
West Germany.
Luthin, C.S. 1984b. Scarlet countdown. Birds
(Lond.) 10:26-32.
Luthin, C.S. 1985a. The bird that's being loved
to death. Naturalist (Leeds) 6(5): 18-21.
Luthin, C.S. 1985b. Captive breeding
programme for the most endangered ibis in the
world. Avic. Mag. 91:117.
Luthin, C.S. 1985c. Suffering scarlet. Naturalist
(Leeds) 6(5):23-27.
Luthin, C.S. 1985d. La ciguena Jabiru. Dumac.
7:6-7.
Luthin, C.S. 1987. Status of and conservation
priorities for the world's stork species. Colonial
Waterbirds 10:181-202.
Luthin, C.S. 1988. Conservation concerns: Indo-
Malayan Realm. Report from the ICBP
Specialist Group on Storks, Ibises and
Spoonbills. Tigerpaper 15:6-7.
Luthin, C.S. 1990. Vorwort [Foreword]. Pp. 9-
16 in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weifistorch: Status und Schutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Luthin, C.S., G.W. Archibald, L. Hartman,
C.M. Miranda, and S. Swengel. 1986. Captive
breeding of endangered cranes, storks, ibises
and spoonbills. Int. Zoo Yb. 24/25:25-39.
Luthin, C.S., and Q. Wang. 1991. Current status
of and conservation concerns for the Oriental
White Stork. Pp. 155-164 in Coulter, M.C., Q.
Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Lutz, A., and C. Meyer. 1908. Hematozoarios
endoglobulares. Rev. Med. (Sao Paulo) 11:177-
183.
Lydekker, R. 1891. On remains of a large stork
from the Allier Miocene. Proc. Zool. Soc. Lond.
1891:476-479.
Lynch, I. 1990. White-faced Ibis at Plum Island,
June 10, 1990. Bird Observer 18(5):290.
Lynes, H. 1925a. On the birds of north and
central Darfur with notes on the West-Central
Kordofan and North Nuba Provinces of British
Sudan. Ibis (ser. 12) 1:71-131, 344-416, 541-
590, 757-797.
Lynes, H. 1925b. L'ornithologie des territoires
du Sous (Maroc du Sud.). Mem. Soc. Sci. Nat.
Maroc 13(1): 1-82.
Lyons, C.M. 1901-1902. Some notes on the birds
of the Lake Eyre District. Emu 1:133-138.
Ma, Y., and L.Jin. 1991. The census of Eastern
White Stork in Heilongjiang Province, China.
Pp. 149-154 in Coulter, M.C., Q. Wang, and
C.S. Luthin (eds), Biology and Conservation of the
Oriental White Stork (Ciconia Boyciana).
Savannah River Ecology Laboratory, Aiken,
South Carolina, USA.
Ma, Y., L.Jin, A.Jin, and C. Fu. 1987. An aerial
survey on Red-crowned Cranes and other rare
waders in the Wuyerhe River Basin of
Heilongjiang Province. Acta Zool. Sin. 833:191.
MacCarone, A.D., and K.C. Parsons. 1988.
Differences in flight patterns of nesting ibises
and egrets. Colonial Waterbirds 11:67—71.
MacDonagh, EJ. 1942. Observaciones sobre la
Bandurria de Invierno. Notas Mus. La Plata
Zool. 7:371-382.
Macdonald, I.A.W., and PJ. Birkenstock. 1980.
Birds of the Hluhluwe-Umfolozi Game
Reserve Complex. Lammergeyer 29:1-56.
MacDonald,J.D. 1957. Contribution to the
Ornithology of Western Africa. William Clowes
and Sons, Ltd, London.
MacDonald, J.D. 1973. Birds of Australia. Reed,
Sydney.
MacDonald, M.A. 1977. Notes on Ciconiiformes
at Cape Coast, Ghana. Nigerian Orn. Soc. Bull.
13:139-144.
Maclnnis, D.G. 1933. Nesting habits of some
East African birds. J. East Afr. Uganda Nat.
Hist. Soc. 11:128-135.
Maclnnis, D.G. 1935. Notes on the birds of
357
Turkana Province. J. East Afr. Nat. Hist. Soc.
Natl. Mus. 12:24-50.
Mackay, H. 1950. The quaint Marabou Stork.
Zoo Life 5:91-92.
Mackey, H.E., Jr. 1984. Agency interaction at the
Savannah River Plant under the endangered species
act. Proceedings of the Fifth D.O.E.
Environmental Protection Conference,
Albuquerque, New Mexico, 5-8 November
1984.
MacKinnon, J.R. 1983. Tanjing Fating National
Park management plan for development. WWF
Proj. 1523 Field Report.
Mackintosh, D.R. 1949. The use of thermal
currents by birds on migration. Ibis 91:55-59.
Mackworth-Praed, C.W., and C.H.B. Grant.
1962. African Handbook of Birds, series II, vol. 1.
Birds of the Southern Third of Africa. Longmans,
Green and Co., London. 688pp.
Mackworth-Praed, C.W., and C.H.B. Grant.
1970. African Handbook of Birds, series III, vol.
1. Birds of West Central and Western Africa.
Longmans, Green and Co., London. 671pp.
Mackworth-Praed, C.W., and C.H.B. Grant.
1973. African Handbook of Birds, series I, vol. 1.
Birds of Eastern and North Eastern Africa.
Longmans, Green and Co., London.
Maclaren, P.I.R. 1952. The sea and creek birds
of Nigeria. Nigerian Field 17:160-174.
Maclean, G.L. 1957. A summary of the birds of
Westminster, O. F. S., and surroundings.
Ostrich 28:217-232.
Maclean, G.L. 1985. Robert's Birds of Southern
Africa, 5th edn. Trustees of the John Voelcker
Bird Book Fund, Cape Town.
Maclean, G.L., R.M. Gous, and T. Bosnian.
1973. Effects of drought on the White Stork in
Natal, South Africa. F0£*/Hwte27:134-136.
Macleod, J.G.R., J. Martin, and CJ. Uys. I960.
Grey-headed Gulls and Spoonbills in the
Bredasdorp area. Ostrich 31:80.
Macmillan, A.T. 1965. Spoonbills in Perthshire
and Moray. Scottish Birds 3:310-311.
Madoc, G.C. 1936. On the nidification of some
Malayan birds. Bull. Raffles Mus. 12:124-133.
Madoc, G.C. 1950. Notes on some birds
occurring in Thailand and Malaya. Part I.
Malay. Nat. J. 5:47-57.
Madoc, G.C. 1956. An Introduction to Malayan
Birds. Malayan Nature Society, Kuala
Lumpur.
Madon, P. 1935. Contribution a 1'etude du
regime des oiseaux aquatiques. Alauda 7:177-
197.
Madsen, S.T. 1990. Black Storks in Nepal and
India. Bull. Oriental Bird Club 11:34-35.
Maehr, D.S., and J. Hintermister. 1982. A
possible White Ibis—Scarlet Ibis hybrid in
Alachua County, Florida. Fla. Field Nat. 10:78.
Mainardi, D. 1962. Immunological data on the
phylogenetic relationships and taxonomic
position of flamingoes (Phoenicopteridae). Ibis
104:426-428.
Mainardi, D. 1963. Immunological distances and
phylogenetic relationships in birds. Proc. Int.
Orn. Congr. 13:103-114.
Majumdar, N. 1989. Past and present wintering
distribution of the White Stork in India and
suggestions for its conservation. Pp. 373-378 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiflstorch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Makowski, H. 1970. Geheimaktion zur Rettung
der letzten Schwarzstorche. Mitteilungsbl.
Schutzgemeinschaft Deutsches Wild (Bonn) I/II:
4-5.
Malchevskii, A.S. 1983. [The White Stork (C.
ciconia) in the Leningrad oblast.] Comm. Baltic.
Commis. Study Bird Migr. 14:119-124.
Malherbe, A. 1855. Faune ornithologique de
1'Algerie. Bull. Soc. Hist. Nat. Moselle 7:5-44.
Malhotra, A.K., and B.M. Arora. 1991. Some
observations on Painted Storks (Mycteria
leucocephalus) at National Zoological Park, New
Delhi. Zoos' Print 6(l):7, 19.
Mallet, M. 1975. The Waldrapp Ibis (Geroniticus
eremita). Jersey WildL Preserv. Trust 12th Annual
Report (1975):26-33.
Mallet, M. 1977. Breeding the Waldrapp Ibis
Geronticus eremita at Jersey Zoo. Int. Zoo Yb.
17:143-145.
Mallinson, JJ.C. 1978. Waldrapp Ibis.
International captive breeding programme for
one of the world's rarest bird species. Zoo Fed.
News 21.
Manakadan, R. 1987. The Black Stork Ciconia
nigra (Linnaeus) in Kurnool District (Andhra
Pradesh)./ Bombay Nat. Hist. Soc. 84:675-676.
Mank, A J. 1963. Die Verbreitung des
Schwarzstorchs in Estland. Thesen Baltischen
Orn. Konf. Tartu 5:120-122.
Mann, C.F. 1989. More notable bird
observations from Brunei, Borneo. Forktail
5:17-22.
Manning, D. and G. Manning. 1971. Some notes
on Glossy Ibis at Reelfoot Lake. Migrant 42:8.
Manry, D.E. 1978a. General biology of the
African ibises. Unpublished report to the
Percy FitzPatrick Institute of African
Ornithology, Univ. of Cape Town,
Rondebosch, South Africa. 23pp.
Manry, D. 1978b. Life with Schaapen Island's
Sacred Ibises— a personal account. Safring
Manry, D.E. 1982. Habitat use by foraging Bald
Ibises Geronticus calvus in western Natal. 6*. Afr.
J. Wildl. Res. 12:85-93.
Manry, D.E. 1983. Ecology of the Bald Ibis
Geronticus calvus and fire in the South African
grassland biome. M.S. Thesis, University of
Cape Town, Cape Town.
Manry, D.E. 1984a. Factors influencing the use
of winter-burnt grassland by foraging Bald
Ibises Geronticus calvus. S. Afr.J. Zool. 19:12-15.
Manry, D.E. 1984b. Pair bonding and promiscuity in
the Bald Ibis (Geronticus calvus). Abstracts,
102nd Stated Meeting of the American
Ornithologists' Union, Lawrence, Kansas, 6-9
August 1984.
Manry, D.E. 1985a. Distribution, abundance,
and conservation of the Bald Ibis (Geronticus
calvus) in southern Africa. Biol. Conserv.
33:351-362.
Manry, D.E. 1985b. Birds of fire. Nat. Hist.
94:38-45.
Manry, D.E. 1985c. Reproductive performance
of the Bald Ibis Geronticus calvus in relation to
rainfall and grass-burning. Ibis 127:159—173.
Manry, D.E. 1986. Ibises in peril. Living Bird Q.
5:24-27.
Manry, D.E. 1990. Living on the edge, Wood
Storks face an uncertain future. Birder's World
4:10-14.
Mansoori, J. 1989. The status of the White Stork
in Iran. Pp. 177-182 in Rheinwald, G., J.
Ogden, and H. Schulz (eds) Weiftstorch: Status
undSchutz, Proc. 1st Int. Stork Conserv. Symp.,
Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Margais, J. 1935. Une colonie d'Ibis chauves
Comatibis eremita (L.) au Maroc oriental. Alauda
7:254-255.
Marcetic, M. 1956. Black Stork, Ciconia nigra, in
Vojvodina. Laray 9-10:153-156.
Marcetic, M. 1957. Roda crna, Ciconia nigra, u
Vojvodini. [The Black Stork, Ciconia nigra, in
Vojvodina.] Larus 9/10:172-175.
Marchant, S. 1958. The birds of the Santa Elena
Peninsula, S.W. Ecuador. Ibis 100:349-387.
Marchant, S. 1963. Migration in Iraq. Ibis
105:369-398.
Marchant, S. and B. Higgins. 1990. Handbook of
Australia, New Zealand and Antarctic Birds, Vol. I,
Part B. Oxford University Press, Oxford,
Auckland and New York.
Marcondes-Machado, L.O., and E.L.A.
Monteiro Filho. 1989. Nota sobre a presenca
dos guaras, Eudocimus ruber (Linne, 1758)
(Threskiornithidae, Aves), no literal de Sao
Paulo. Alerta para sua protecao. Ciencia e
Cultura 41:1213-1214.
Marcondes-Machado, L.O., and E.L.A.
Monteiro Filho. 1990. The Scarlet Ibis
Eudocimus ruber in southeastern Brazil. Bull. Br.
Orn. Club 110:123-126.
Mari, M.E.L. 1984. Contribucion al estudio de la
alimentacion de la Ciguena Blanca, Ciconia c.
ciconia (L.) en Espana. Thesis Doctoral,
Universidad Complutense de Madrid.
Marian, M. 1970. Der Bestand des WeiBstorchs
(C. ciconia) in Ungarn 1963. Vogelwarte 25:255-
257.
Marion, L., and P. Marion. 1982. [The Spoonbill
(Platalea leucorodia) nests at the Lac de Grand-
Lieu, France.] Alauda 50:241-249.
Marie, J.G. v., and K.H. Voous. 1988. The Birds
of Sumatra: an Annotated Check-list Br. Ornithol.
Union 10, Tring.
Marno, E. 1874. Die Stoerche des Nilgebietes.
Zool. Gart. 15:48-55.
Marshall, B.E. 1982. A possible example of tool
usage by the Marabou Stork. Ostrich 53:181.
Martens, J. 1966. Brutvorkommen und
Zugverhalten des Weiflstorchs (Ciconia ciconia)
in Griechenland. Vogelwarte 23:191-208.
Martin, E., R. Martin, and J. Robinson. 1962.
European stork Ciconia ciconia breeding in the
Bredasdorp district. Ostrich 33:26-27.
Martin, J., E. Martin, and R. Martin. 1976.
Notes from western Calvinia. Bokmakierie
28:20-23.
Martin, RJ. 1971. Hadedah Bostrychia hagedash.
Ostrich 42:137.
Martin, RJ. 1972. Hadeda nesting in the
southern entrance of the Seven-Weeks Poort.
Ostrich 43:185-186.
Maslowski, K.H., and P. Koch. 1940.
Adventures with the Roseate Spoonbill. Nat.
Hist. 45:108-111.
Masterson, A.N.B. 1971. Three weeks in
Botswana. Honey guide 65:11—18.
Masterson, A.N.B. 1973. Openbill Storks
breeding in Rhodesia. Honeyguide 76:39.
Mathena, C., J. Hickman, J. Hill, R. Mercer, C.
Berry, B. Berry, N. Berry, N. Cherry, and P.
Flynn. 1984. Birds of Dayton. Landfall Press,
Dayton, Ohio.
Mathews, G.M. 1910. On birds of north-west
Australia. Emu 10:103-110.
Mathews, G.M. 1913. On the generic names Ibis
Lacepede, and Egateus Billberg. Auk 30:92-95.
Mathews, NJ.C. 1979. Observations of the
Shoebill in the Okavango Swamp. Ostrich
50:185.
Mathiasson, S. 1963. Visible diurnal migration
in the Sudan. Proc. Int. Orn. Congr. 13:430-435.
Matthews, G.V.T. 1990. Managing Waterfowl
Populations. Proc. IWRB Symp., Astrakhan
1989. IWRB Special Publication 12. IWRB,
Slimbridge, U.K.
Mattocks, J. 1948. Possible observation of Wood
Ibis near High Point, N.C. Chat 12:65-66.
Maxwell, G.R. II, and H.W. Kale II. 1977.
Maintenance and anti-insect behavior of six
species of ciconiiform birds in south Florida.
Condor 79:51-55.
Mayaud, N. 1950a. Longevite. Pp. 536-538 in
Grasse, P.-P. (ed.), Traite de Zoologie (Oiseaux,
Tome 15). Masson and Cie, Paris.
Mayaud, N. 1950b. Alimentation. Pp. 654-688 in
Grasse, P.-P. (ed.) Traite de Zoologie (Oiseaux,
Tome 15). Masson and Cie, Paris.
Mayaud, N. 1982. Les oiseaux du Nord-Ouest de
1'Afrique: notes complementaries. Alauda
50:114-145.
358
Maynard, C J. 1881. The Birds of Eastern North
America. C J. Maynard, Newtonville,
Massachusetts.
Mayr, E. 1938. Notes on a collection of birds
from south Borneo. Bull. Raffles Mus. 14:5-46.
Mayr, E. 1989. A new classification of the living
birds of the world. Auk 106:508-512.
Mayr, E., and G. Cottrell. 1979. Checklist of Birds
of the World, vol. 1. Museum of Comparative
Zoology, Cambridge, Massachusetts.
Mayr, E., and L.L. Short. 1970. Species taxa of
North American birds. Publ. Nuttall Orn. Club
9:1-127.
McAllan, I.A.W., and M.D. Bruce. 1989. The
Birds of New South Wales: a Working List. Biocon
Research Group, Turramurra, NSW. 103pp.
McAllister, V. 1977. Possible predation by
Tawny Eagle on White Stork. Honeyguide
89:47.
McAlpine, D.F., J. Finne, S. Makepeace, S.
Gilliland, and M. Phinney. 1989. First nesting
of the Glossy Ibis, Plegadisfalcinellus, in
Canada. Can. Field-Nat. 102:536-537.
McAtee, W.L. 1913. Two ornithological fables
from Louisiana. Auk 30:282.
McCann, C. 1930. Nidification of storks. J.
Bombay Nat. Hist. Soc. 34:579-581.
McCarley, H., and K.W. Haller. 1984. Wood
Stork in Lake Texoma area. Bull. Okla. Orn.
Soc. 17:14-15.
McClure, H.E. 1974. Migration and Survival of the
Birds of Asia. U.S. Army component SEATO
medical research laboratory Bangkok,
Thailand. 476pp.
McClure, H.E., and P. Kwanyuen. 1973. The
avifaunal complex of an Open-billed Stork
colony (Anastomus oscitans) in Thailand. Nat.
Hist. Bull. Siam Soc. 25:133-178.
McConnell, E.S., and J.P. McConnell. 1974.
Jabiru Stork in Oklahoma. Bull. Oklahoma Orn.
Soc. 7:9-12.
McCoy, J J. 1963. The fossil avifauna of
Itchtucknee River, Florida. Auk 80:335-351.
McCulloch, E.M. 1974. Sexual difference in eye-
colour of Jabiru, Xenorhynchus asiaticus. Aust.
BirdWatcher 5:187-188.
McDaniel, B., E. Bogusch, and S. McDaniel.
1963. A unique behavior pattern and
observations of Roseate Spoonbills (Ajaia ajaja)
in Kleberg County, Texas. Tex. J. Sci. 15:354-
356.
McGill, A.R. 1944. The Glossy Ibis. Emu 43:280.
McGill, A.R. 1958. Branch report. Emu 58:44-
45.
McKilligan, N.G. 1975. Breeding and movement
of the Straw-necked Ibis in Australia. Emu
75:199-212.
McLachlan, G.R. 1963. European stork Ciconia
ciconia ringed as nestling in South Africa
recovered in Northern Rhodesia. Ostrich 34:48.
McLachlan, G.R. 1966. The first ten years of
ringing in South Africa. Ostrich suppl. 6:255-
263.
McLachan, G.R., and R. Liversidge. 1970.
Roberts' Birds of South Africa, 3rd edn. Trustees
John Voelcker Book Fund, Cape Town. 634pp.
MacLachan, G.R., and R. Liversidge. 1978.
Robert's Birds of South Africa, 4th edn. Trustees
John Voelcker Book Fund, Cape Town.
McLean, S. 1986. Saddlebilled Stork breeding at
St. Lucia. Lammergeyer 5*7:55.
McNabb, G.C. Jr. 1978. White Ibis near
Rosman, N.C. Chat 42:81.
McNeil, R., B. Limoges, and J. R. Rodriguez S.
1990. El Corocoro Rojo (Eudocimus ruber) y
otras aves acuaticas coloniales de las lagunas,
cienagas, y salinas de la costa centro-oriental
de Venezuela. Pp. 28-45 in Frederick, P.C.,
L.G. Morales, A.L. Spaans, and C.S. Luthin
(eds), The Scarlet Ibis (Eudocimus ruber):
Status, Conservation, and Recent Research, Proc. 1st
International Scarlet Ibis Conservation
 Workshop, Caracas, Venezuela. IWRB
Special Publication 11, Slimbridge, Glos, UK.
McShane, T.O., and E. McShane-Caluzi. 1988.
The habitats, birds and mammals of Vwaza
Marsh Game Reserve, Malawi. Nyala 12:39-
66.
Meade-Waldo, E.G.B. 1903. Bird-notes from
Morocco and the Great Atlas. Ibis 8:196-214.
Meade-Waldo, E.G.B. 1905. A trip to the forest
of Marmora, Morocco. Ibis 8:161-164.
Mebs, T. 1969. Der Bestand des WeiBstorchs
(Ciconia ciconia) in Franken 1965-1967 im
Vergleich mit den Ergebnissen fruherer
Zahlungen. Anz. Orn. Ges, Bayern. 8:463^72.
Medway, Lord, and D.R. Wells. 1964. Bird
report: 1963. Malay. Nat.J. 18:133-167.
Medway, Lord, and D.R. Wells. 1971. Diversity
and density of birds and mammals at Kuala
Lompat, Pahang. Malay. Nat.J. 24:238-247.
Medway, Lord, and D.R. Wells. 1976. The Birds
of the Malay Penninsula, vol. 5. H.F. and G.
Witherby, London.
Meer-Mohr, J.C. v.d. 1923. lets over een bango.
Trap. Natuur 12:62-63.
Mees, G.F. 1982. Bird records from the
Moluccas. Zoo/. Meded. (Leiden) 56:94-111.
Meinertzhagen, R. 1920. Some preliminary
remarks on the altitude of the migratory flight
of birds, with special reference to the
Palearctic region. Ibis (ser. 11) 2:920-936.
Meinertzhagen, R. 1930. Nicoll's Birds of Egypt.
Hugh Rees Ltd., London.
Meinertzhagen, R. 1937. Some notes on the birds
of Kenya Colony, with special reference to
Mount Kenya. Ibis (ser. 4) 1:731-760.
Meinertzhagen, R. 1940. Autumn in central
Morocco. Ibis 14(4): 106-136, 187-234.
Meinertzhagen, R. 1951. Some relationships
between African, Oriental, and Palearctic
genera and species, with a review of the genus
Monticola. Ibis 93:443-459.
Meinertzhagen, R. 1954. Birds of Arabia. Oliver
and Boyd, London.
Meinertzhagen, R. 1955. The speed and altitude
of bird flight (with notes on other animals).
/My 97:81-117.
Meinertzhagen, R. 1959. Pirates and Predators.
Oliver and Boyd, London.
Meise, W. (ed.). 1960-1963. Handbuech der
Oologie. Akademie-Verlag, Berlin.
Meitz, P. 1979a. Marokkom, Heimat des
Waldrapp. Gefaderte Welt 103:78-80, 94-97.
Meitz, P. 1979b. Noch einmal in Marokko-des
Waldrapps wegen! Gefaderte Welt 103:193-194.
Meitz, P. 1980. Nochmals: vom Waldrapp in
Marokko. Gefiederte Welt 104:119.
Mell, R. 1951. DerStorch. Die Neue Brehm-
Buecherei, Heft 35. Wittenberg/Lutherstadt.
Mellen, J.W., and N.O. Dronen, Jr. 1987. Some
digenetic trematodes from a Wood Ibis
Mycteria americana, including a new species of
Ascocotyle. Trans. Amer. Microsc. Soc. 106:269-
272.
Mendelssohn, H. 1975a. The White Stork
(Ciconia ciconia) in Israel. Vogelwarte 28:\23-
131.
Mendelssohn, H. 1975b. Report on the status of
some bird species in Israel in 1974. Bull. ICBP
12:265-270.
Menon, G.K., R.V. Shah, and M.B. Jani. 1979.
Lipoid secretion by epidermis of bare skin
from the head of the Indian White Ibis
Threskiornis melanocephala. J. Yamashina Inst. Orn.
11:128-131.
Menspira, M.A. 1895. [Russian Birds.] Moscow.
Merriam, C.H. 1890. Results of a biological survey of
the San Francisco mountain region and desert of Little
Colorado, Arizona. U.S. Dept. Agric. N. Am.
Fauna 3. Washington DC.
Merrill, J.C. 1879. A nesting place of the White-
faced Glossy Ibis (Plegadis guarauna). Oologist
Merz, A. 1980. Marabou Stork kills Lesser
Flamingo. Bull. East Afr. Nat. Hist. Soc. Nov/
Dec:97-98.
Metzen, W.D. 1977. Second nesting of Wood
Storks in Georgia. Oriole 42:30-31.
Mey, E. 1988. Herbstdaten zur mongolischen
Avifauna. Mitt. Zool. Mus. Berl. 64 (suppl.):
Ann. Orn. 12:79-128.
Meybohm, E. 1970. 22 jahriger WeiBstorch (C.
ciconia) brudtet und zieht drei Junge auf.
Vogelwarte 25:267.
Meybohm, E., and G. Dahms. 1975. Uber
Altersaufbau, Reifealter und Ansiedlung biem
WeiBstorch (Ciconia ciconia) im Nordsee-
Kuestenbereich. Vogelwarte 28(1):44-61.
Meyer, A.B., and L.W. Wiglesworth. 1898. The
Birds of Celebes', vol. 2. Friedlander, Berlin.
Meyer de Schauensee, R. 1946. On Siamese
birds. Proc. Acad. Nat. Sci. Phila. 98:1-82.
Meyer de Schauensee, R. 1948. The birds of the
Republic of Colombia. Pt. 1. Caldasia 5:251-
379.
Meyer de Schauensee, R. 1964. The Birds of
Colombia. Livingston Publ., Narbeth, Pa.
427PP.
Meyer de Schauensee, R. 1966. The Species of
Birds of South America and Their Distribution.
Livingston Publ. Co., Narbeth, Pennsylvania.
577pp.
Meyer de Schauensee, R. 1970. A Guide to the
Birds of South America. Oliver and Boyd,
Edinburgh.
Meyer de Schauensee, R. 1984. The Birds of
China. Oxford University Press, Oxford.
Meyer de Schauensee, R., and W.H. Phelps, Jr.
1978. A Guide to the Birds of Venezuela. Princeton
University Press, Princeton, New Jersey.
Meyerriecks, AJ. 1959. Foot-stirring feeding
behavior in herons. Wilson Bull. 71:153-158.
Meyers, J.M. 1980. First White Ibis record for
Athens, Georgia. Oriole 45:41-42.
Meyers, J.M. 1984. Wood Storks of the Birdsville
Colony and swamps of the Savannah River Plant.
SREL-15/UC-66e. Savannah River Ecology
Laboratory, Aiken, SC.
Michalega, P. 1979. Nove vyslcyty capu cernych
(Ciconia nigra L.) v oblastikrusnych hor a v
CHICO Ceske stredohori. Fauna Bohemiae
Septentrionalis 4:11-12.
Michelmore, F., and W.L.R. Oliver. 1982.
Hand-rearing and development of Bare-faced
Ibis chicks (Geronticus eremitd) at the Jersey
Wildlife Preservation Trust: with comparative
observations of parent-rearing behavior. Dodo
19:51-69.
Michev, T.M., and L.T. Stojanova. 1986.
[Nesting distribution and numbers of the
White Stork Cicodnia ciconia (L. 1758).]
Ekologiya (Sofia) 18:17-26.
Michev, T., and L. Profirov. 1989. Der
Herbstzug des WeiBstorchs entlang der
bulgarischen Schwarzmeerkuste. Pp. 281-296
in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weifistorch: Status und Schutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Michev, T.M., N.P. Dilchev, and S;D.
Simeonov. 1985. [Dynamics of numbers and
nest biology of the White Stork Ciconia ciconia
(L.) within the region of Sofia.] Ekologiya
(Sofia) 17:49-55.
Michev, T., T. Petrov, and L. Profirov. 1989.
Status, breeding, distribution, numbers and
conservation of the White Stork in Bulgaria.
Pp. 137-144 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weijtstorch: Status undSchutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Middlemiss, E.HJ. 1955. Nest of Glossy Ibis at
Rondevlei. Ostrich 26:137-138.
359
Mikolas, K. 1929. Ciconia alba und die rote
Farbe. Aquila 34-35:446.
Miles, H.M. 1951. The present status of the
White Stork in S. Rhodesia. Ostrich 22:195-
196.
Miller, I.D. 1990. The Rare Birds of Tula's Land.
Kommunar, Tula, USSR.
Miller, I.G. 1989. Ciconia ciconia in Tula's region:
last and recent. Report on the First Meeting of
the USSR Stork Research Group, July 1989,
Tallinn, USSR.
Miller, J.C. 1973. Second nesting of herons,
egrets and ibis. Cassinia 54:29.
Miller, J.C. 1982a. White Ibis in Delaware
County, Pennsylvania. Cassinia 59:82.
Miller, J.C. 1982b. White Ibis in Philadelphia
County, Pennsylvania. Cassinia 59:82.
Miller, L. 1932. The Pleistocene storks of
California. Condor 34:212-216.
Miller, L. 1944. Some Pliocene birds from
Oregon and Idaho. Condor 46:25-32.
Miller, L. and R. Bowman. 1958. (P. gracilis)
description from Pleistocene of Texas. Los Ang.
Cty Mus. Contrib. Sci. 20.
Miller, L.H. 1910. Wading birds from the
Quaternary asphalt beds of Rancheo La Brea.
Univ. Calif. Publ. Geol. 5:439-448.
Miller, L.M. 1976. Expansion dynamics and
breeding biology of the Glossy Ibis, Plegadis
falcinellus. M.S. Thesis, Rutgers University,
New Brunswick, New Jersey.
Miller, L.M., and J. Burger. 1978. Factors
affecting the nesting success of the Glossy Ibis.
4M* 95:353-361.
Miller, W.T. 1947. Birds at Home. Camera Studies
of Birds of South Africa. Maskew Miller, Cape
Town, South Africa.
Mills, E.L. 1967. Bird records from southwestern
Ecuador. Ibis 109:534-538.
Mills, M.G.L. 1976. Glossy Ibis Plegadis
falcinellus. Ostrich 47r:216.
Milne-Edwards, A. 1867-1871. Recherches
Anatomiques et Paleontologiques pour Servir a
I' Histoire des Oiseaux Fossiles de la France, 4 vols.
Masson et Fils, Paris.
Milne-Edwards, A., and A. Grandidier. 1879-
1885. Oiseaux (Supplement vor L. Lavanden,
1937) In A.U.G. Grandidier Historie Physique,
Naturelle et Politique de Madagascar, vol. XII.
Paris.
Milon, P., et al. 1973. Faune de Madagascar,
Oiseaux [in Mayr and Cottrell, 1979:254].
Milstein, P. le S. 1965. Uber BHC-vergiftete
Heuschrecken als Nahrung des WeiBstorchs in
Sudafrika. Vogelwarte 23:117-121.
Milstein, P. le S. 1966. Preliminary observations
of White Storks feeding on poisoned brown
Locusts. Ostrich suppl. 6:197-215.
Milstein, P. le S. 1973. Buttons and Bald Ibises.
Bokmakierie 25:57-60.
Milstein, P. le S. 1974. More Bald Ibis buttons.
Bokmakierie 26:88.
Milstein, P. le S. 1975. The biology of
Barberspan, with special reference to the
avifauna. Ostrich suppl. 10:1-74.
Milstein, P., and WA. Siegfried. 1970.
Transvaal status of the Bald Ibis. Bokmakierie
22:36-39.
Milstein, P., and S.W. Wolff. 1973. Status and
conservation of the Bald Ibis in the Transvaal.
J. S. Afr. Wildl. Manag. Assoc. 3:79-83.
Milton, G.R., and A. Marhadi. 1985. The bird
life of the nature reserve Pulau Dua. Kukila
2:32-41.
Mitchell, C.A. 1931. Roseate Spoonbills in
Florida. Auk 48:259.
Mitchell, C.W. 1983. White Ibis observed on
Lake Champlain in Clinton County. Kingbird
33:34-35.
Mitchell, J. 1972. Spoonbill in Durbartonshire
and West Stirlingshire. West. Nat. 1:114-115.
Mitchell, P.C. 1894. On the perforated flexor
 muscles in some birds. Proc. ZooL Soc. Lond.
1894:495-498.
Mitchell, P.C. 1895. On the proventricular
crypts of Pseudotantalus ibis. Proc. ZooL Soc. Lond.
1895:271-273.
Mitchell, P.C. 1896. On the intestinal tract of
birds. Proc. ZooL Soc. Lond. 1896:136-159.
Mitchell, P.C. 1911. On longevity and relative
viability in mammals and birds; with a note on
the theory of longevity. Proc. ZooL Soc. Lond.
1911:425-548.
Mitchell, P.C. 1913. Observations on the
anatomy of the Shoe-bill (Balaeniceps rex) and
allied birds. Proc. ZooL Soc. Lond. 1913:644-
703.
Mitchell, P.C. 1915. The stomach and intestines
of the Open-bill. Proc. ZooL Soc. Lond.
1915:153.
Mizuno, K. 1934. A Distributional List of the
Manchurian Birds. Nagahisa Varoda, Tokyo.
Mlingwa, C. 1989. Notes on Marabou Storks
Leptoptilos crumeniferus at Shinyanga, Tanzania.
E. Afr. Nat. Hist. Soc. Bull. 19:34.
Mock, D.W. 1978a. Behavior and evolution of
wading birds: introductory remarks. Pp. 3-6 in
Sprunt, A., J.C. Ogden, and S. Winckler (eds),
Wading Birds. Research report No. 7, National
Audubon Society, New York.
Mock, D.W. 1978b. The comparative approach
to wading bird behavior. Pp. 17-25 in Sprunt,
A., J.C. Ogden, and S. Winckler (eds), Wading
Birds. Research report 7, National Audubon
Society, New York.
Mohammed, S. 1985. The Caroni Swamp.
Naturalist 6:1-81.
Molina, A. 1782. Saggio Storia Nat. Chili.
Mollendorf, O.F. v. 1877. The vertebrata of the
Province ofChihli. II. birds. J. N. Asiatic Soc.
11:76-102.
Moller, W. 1978. Auf den Spuren von Konig
Walkopf. Sielmanns Tierwelt 8:36-44.
Moller, W. 1979. Africa's amazing Shoebill Stork
is a whale of a bird. Wildlife (Lond.) 21:38-39.
Moller, W. 1980. Freilandbeobachtungen am
Schuhschnabel—Balaeniceps rex. Diplomarbeit
des Zool. Inst. der TU Braunschweig.
Moller, W. 1982a. Beobachtungen zum
Nahrungserwerb des Schuhschnabels
(Balaeniceps rex). [Observations on the feeding
behavior of the Shoebill (Balaeniceps rex).]J.
Orn. 123:19-28.
Moller, W. 1982b. Bei den Schuhschnabeln in
den Sumpfen Ugandas. Zoofreunde Hannover
45:6-10.
Moltoni, E., and G. Ruscone. 1942. Gli uccelli dell
'Africa Orientate. Italiana. 2, Milan.
Monroe, B.L. 1938. Some rarer birds at the Falls
of the Ohio. Auk 55:678.
Monroe, B.L., Jr. 1968. A Distributional Survey
of the Birds of Hondurus. A.O.U.
Ornithological Monograph 7.
Montagu, G. 1818. Some remarks on the natural
history of the Black Stork for the first time
captured in Great Britain. Trans. Linn. Soc.
Lond. 12:19-23.
Montfort, N. 1965. Ornithologische
Beobachtungen imjemen.j. Orn. 106:333-
339.
Montgomery, K. 1988(1990). Roseate Spoonbill
seen in east-central Georgia. Oriole 53:26-27.
Moore, HJ. and C. Boswell. 1956. Field
observations on the birds of Iraq. Iraq Nat.
Hist. Mus. Publ. 9:1-109; 10:110-213; 12:214-
299.
Moore, J.C. 1953. Glossy Ibis in the Everglades
National Park. Everglades Nat. Hist. 1:25-28.
Morales, G. 1990. Conservacion de las aves
zancudas en los llanos de Venezuela. Pp. 77-
84 in Frederick, P.C., L.G. Morales, A.L.
Spaans, and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research. Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Morales, G., and N. Leon. 1983. Las aves
Pescadores de los modulos de Mantecal. Natura
73:22-27,
Morales, G., J. Pinowski, J. Pacheco, M. Madriz,
and F. Gomez. 1981. Densidades
poblacionales flugo de energia y habites
alimentaries de las aves ictiofages de
losmodulos de puve, Venezuela. Acta Biol.
Venez. 11:1-45.
Moreau, R.E. 1928. Some further notes from the
Egyptian deserts. Ibis (ser. 12) 4(3):453-475.
Moreau, R.E. 1930. The birds of ancient Egypt.
Pp. 58-77 in Meinertzhagen, R. (ed.), Nicoll's
Birds of Egypt, vol. 1. Hugh Rees, London.
Moreau, R.E. 1942. The nesting of African birds
in association with other living things. Ibis
(ser. 14) 6:240-263.
Moreau, R.E. 1950. The breeding seasons of
African birds-1. land birds. Ibis 92:223-267.
Moreau, R.E. 1953. Migration in the
Mediterranean area. Ibis 95:329-364.
Moreau, R.E. 1959. Les problemes de la
migration a travers le Maroc. Alauda 27:81-96.
Moreau, R.E. 1966a. The mutability of the
African avifaunal scene. Ostrich suppl. 6:453-
459.
Moreau, R.E. 1966b. The Bird Faunas of Africa and
its Islands. Academic Press, New York.
Moreau, R.E. 1967. Water-birds over the
Sahara. Ibis 109:232-259.
Moreau, R.E. 1970. Changes in Africa as a
wintering area for Palaearctic birds. Bird Study
17:95-103.
Moreau, R.E. 1972. The Palaearctic African Bird
Migration Systems. Academic Press, London.
Moreau, R.E., and W.L. Sclater. 1938. The
avifauna of the mountains along the Rift
Valley in North Central Tanganyika Territory
(Mbulu District). II. Ibis (ser. 14) 2(l):l-32.
Morel, G., and M.-Y. Morel. 1961. Une
heronniere mixte sur le Bas-Senegal. Alauda
29:99-117.
Morel, G., and M.-Y. Morel. 1962. La
reproduction des oiseaux dans une region
semi-aride: La Vallee du Senegal. Alauda
30:161-203,241-269.
Morel, G., and F. Roux. 1966. Les migrateurs
palearctiques au Senegal. La Terre et la Vie
113:19-72, 143-176.
Morgan-Davies, A.M. 1964. Notes on some
breeding birds of Lake Manyara National
Park. Tanganyika Notes Rec. 62:96-104.
Mori, T. 1938. [On the breeding ground and
habits ofdconia nigra.] Tori 10:127-129.
Moriearty, P.L., D.E. Pomeroy, and B. Wanjala.
1972. Parasites of the Marabou Stork
(Leptoptilos crumeniferus (Lesson)) in Queen
Elizabeth National Park, Uganda. E. Afr.
Wildl.J. 10:311-314.
Moriguchi, K. 1977. [A record of(Ciconia ciconia)
found on Shiodome River near Hakodate City,
Hokkaido.] Ton 26(4): 129.
Morioka, H., and C. Yano. 1990. A record of the
Milky Stork in Thailand. Jap.J. Orn. 38:149.
Morris, A.K. 1973. White ibis eats snake. Emu
73:73.
Morris, F.T. 1978. Feeding association between
Little Egret and Sacred Ibis. Emu 78:164.
Morris, A.K. 1979. Saddlebill fishing methods.
Honeyguide 98:33.
Morris, G.E. 1989. Further (1988) sight records
of birds at Nam Cat Tien. Garrulax 6:2.
Morris, P.J., and J.P. Weigel. 1986. Mandibular
fracture repair in a Marabou Stork (Leptoptilos
crumeniferus) with a methacrylate prosthesis.
Proc. Annu. Meet. Assoc. Avian Vet. 1986:303-
305.
Morrison, M.L., R.D. Slack, and E. Shanley, Jr.
1978. Interspecific association of Olivaceous
360
Cormorants and Roseate Spoonbills. Southwest
Nat. 23:681-683.
Morrison, R.G., R.K. Ross, P. Canivari, Z.
Antas, B. dejong, B. Ramdial, F. Espinosa,
M.M. Teperino, and J. Mago de Perez. 1985.
Aerial surveys of shorebirds and other wildlife
in South America: some preliminary results.
Can. Wildl. Sew. Prog. Notes 148.
Morse, C.M. 1965. Jabiru in north-west New
South Wales. Emu 64:324.
Moser, D. 1971. A lament for some doomed
companions. Life 22 January 1971.
Mott, L.T. 1965. Water birds of the Gran Chaco
of Paraguay. Game Bird Breeders Gaz. 14:14—15.
Mountfort, G. 1958. Portrait of a Wilderness.
Hutchinson, London.
Mountfort, G. 1963. The Wild Danube: Portrait of a
River. Hough ton Mifflin, Boston.
Mountfort, G., and I J. Ferguson-Lees. 1961.
Observations on the birds of Bulgaria. Ibis
103a:443-471.
Mountfort, G., and D. Poore. 1968. Report on
the second World Wildlife Fund expedition to
Pakistan. Unpublished report on World
Wildlife Fund Project 311. WWF, Reigate,
UK.
Mouquet, A., and C. Truche. 1924. Gangrene
des extremites inferieures chez les oiseaux.
Bull. Mus. Nat. d'Hist. Nat. Paris 30:443-449.
Mourer-Charvire, C., and F. Moutou. 1987.
Decouverte d'une forme recemment eteinte
d'ibis endemique insulaire de 1'ile de la
Reunion Borbonibis latipes n. gen. n. sp. C. R.
Acad. Sci. Paris (ser. II) 305:419-423.
Moynihan, M. 1970. Control, suppression,
decay, disappearance and replacement of
displays./ Theor. Biol. 29:85-112.
Mracovsky, F.H. 1976. Dalsi skalni hnizda capa
cerneho (Ciconia nigra L.) na Morave. Zpravy
MOS 18(34):! 17-121.
Mrugasiewicz, A. 1972. White Stork, Ciconia
ciconia (L.) over the district of Milicz in the
years 1959-1968. Acta Orn. (Warsaw) 13:243-
278.
Mrugasiewicz, A., and J. Witkowski. 1962. An
ornithological sketch of the Barycz Valley in
Poland. British Birds 55:245-272.
Muchmore, D. 1973. White-faced Ibis. Wyoming
Wildl. 37:24-25.
Mukherjee, A.J. 1974. Food habits of water-birds
of the Sunderban, 24 Parganas District, West
Bengal, India—IV. J. Bombay Nat. Hist. Soc.
71:188-200.
Mukhopadhyay, A. 1980. Some observations on
the biology of the Openbill Stork, Anastomus
oscitans (Boddaert) in southern Bengal, y.
Bombay Nat. Hist. Soc. 77:133-137.
Muller, C.Y. 1983. Die Bedeutung von
Altschilfbestanden fur die Loffler und Reiher
am Neusiedlersee. Egretta 26:43—46.
Muller, C.Y. 1984. Bestandsentwicklung und
Zugverhalten der Loffler (Platalea leucorodia L.)
im osterreichisch-ungarischen Raum. Egretta
27:45-67.
Muller, C.Y. 1987. Feeding-sites and resting-
places of the Spoonbill Platalea leucorodia at
Neusiedlersee (Austria). Orn. Beob. 84:237-
245.
Muller, C.Y. 1987. [Contribution to
reproduction and rearing of the Spoonbill
(Platalea leucorodia L.) in the region of
Neusiedlersee]. Egretta 30:13-23.
Miiller-Scheessel, L. 1964. Zum Brutvorkommen
des Schwarzstorches in Niedersachsen. Beitr.
Naturkd. Niedersachsens 17:60-62.
Muller-Scheessel, L. 1965. Ueber
Brutvorkommen des Schwarzstorches in
Niedersachsen im Jahre 1964. Beitr. Naturkd.
Niedersachsens 18:19-20.
Mumford, R.E., and L.E. Lehman. 1969. Glossy
ibis taken in Indiana. Wilson Bull. 81:463-464.
Mundy, PJ. 1985. What brings the African
Marabou on? Honey guide 31:186.
Mundy, PJ. 1988. Marabou Storks breeding at
Kadoma. Honeyguide 34:66-69.
Mundy, P. 1989. A roost of egrets and ibises at
Lake Mcllwaine. Honeyguide 35:112.
Mundy, PJ., and A.W. Cook. 1974. The birds of
Sokoto. Part 3. Breeding data. Bull. Niger. Orn.
Soc. 10:1-28. .
Mundy, P., I.C. Cannell, and A. Williams. 1988.
Marabou Storks breeding at Kasoma.
Honeyguide 34:66-69.
Munetchika, I., M. Morita, and S. Watanabe.
1976. Studies on the serological relationships
among some storks (Ciconiidae). Tori 25:109—
113.
Munzinger, S. 1985. Weiss-Storch (Ciconia
ciconia) attackiert Schwarzstorch (Ciconia
nigra). Orn. Mitt. 37:20.
Murata, K. 1984. Preparation and attachment of
a prosthetic bill in a White Stork. J.Japan. Vet.
Med. Assoc. 37:805-808.
Murata, K. 1988. Sex determination in the
Eastern White Stork, Ciconia c. boyciana, by bill
measurements and discriminant analysis. J.
Jpn. Assoc. Zoos Aquariums 30:43-47.
Murdock, N.A. 1987. Creation of artificial
foraging habitat for Wood Storks. Endang.
Species Tech. Bull. 12(1):4, 16.
Murphey, E.E. 1937.. Observations on the Bird Life
of the Middle Savannah Valley, 1890-1937.
Contributions from the Charleston Museum
IX.
Murphey, E.E. 1939. Concerning the Wood Ibis.
P. 16 in Wings at Dusk and Other Poems.
Longmans, Green, and Co., New York.
Murphy, R.C. 1925. Bird Islands of Peru. New
York.
Museo Argentine de Ciencias Naturales. 1935.
Lista sistematica de las aves argentinas.
Hornero 6:151-196.
Musselman, T.E. 1968. Chuck-will's-widow and
Wood Ibis in central Illinois. Wilson Bull.
80:230.
Muzinic, J. 1989. Distribution and status of the
White Stork in Croatia (Yugoslavia) in 1984.
Pp. 121-124 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp.., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Myong, J.S. 1967. Is the White Stork really
extinct? Korean Nat. 1967:2-3.
Nacinovic, J.B., and D.M. Teixeira. 1989. As
aves de Fernando de Noronha: uma lista
sistematica anotada. Rev. Brasil Biol. 49:709-
729.
Naggiar, M. 1975. The White Ibis. Florida
Wildlife 2<M3V90-21.
Nagy, I. 1955. [Biological observations on a pair
of White Storks with special consideration for
the feeding of the young.] Aquila, 59—62:145-
159.
Nakayama, T. 1980. [Handraising the Scarlet
Ibis.] Animals and Zoos 32:152-154.
Nakhasathien, S. 1987. The discovery of Storm's
Stork Ciconia stormi in Thailand. Forktail 3:43-
49.
Nash, S.V., and A.D. Nash. 1985. A checklist of
the forest and forest edge birds of the Padang-
Sugihan Wildlife Reserve, South Sumatra.
Kukila 2:5 1-59.
Nash, S.V., and A.D. Nash. 1988. An annotated
checklist of the birds of Tanjung Puting
National Park, Central Kalimantan. Kukila
3:93-116.
Narosky, T. 1978. Aves Argentinas. Assoc. Ornit.
del Plata, Buenas Aires.
Naumann,J.A. 1838. Naturgeschichte der Vogel
DeutschlandS) vol. 9. Fleischer, Leipzig.
Naumburg, E.M. 1930. The birds of Matto
Grosso, Brazil. Bull. Am. Mus. Nat. Hist. 60:1-
432.
de Naurois, R. 1959. Premieres recherches sur
1'avifaune des iles du Bane d'Arguin
(Mauritanie). Alauda 27:241-308.
de Naurois, R. 1966. Colonies reproductrices de
Spatules africaines, Ibis sacres et lorides, dans
L'Archipel des bijagos (Guinee Portugaise).
Alauda 34:257-278.
de Naurois, R. 1969. Peuplements et cycles de
reproduction des oiseaux de la cote occidentale
d'Afrique. Mem. Mus. Natl Hist. Nat. Ser. A
Zoo/. 56:1-293.
de Naurois, R. 1973. Les ibis des iles de Sao
Tome et du Prince: leur place dans le groupe
des Bostrychia (=Lampribis). Archives. Mus.
Bocage(ser. 2)4:157-173.
de Naurois, R., and F. Roux. 1974. Precisions
concernant la morphologic, les affinites et la
position systematique de quelques oiseaux du
Bane d'Arguin (Mauritanie). Oiseau Rev. Fr.
Orn. 44:72-84.
Nazarov, Y.N., and V.I. Labzyuk. 1975. On the
avifauna of South Primorye. Rep. Biol. Inst.
29:268-276.
Neame, G.B. 1968a. Glossy Ibis, Plegadis
falcinellus, distribution. Ostrich 39:271.
Neame, G.B. 1968b. Spoonbill breeding in the
Eastern Cape. Ostrich 39:265.
Neginhal, S.G. 1977. Discovery of a pelicanry in
Karnataka.J. Bombay Nat. Hist. Soc. 74:169-
170.
Neginhal, S.G. 1982. The birds of
Ranganathittu.y. Bombay Nat. Hist. Soc.
79:581-593.
Nehkorn, A. 1876. Afrikanischer Nimmersatt
und wollhalsiger Storch. Zoo/. Gart. 17:414.
Neis, K. 1979. Der Heilige Ibis. Zoologischen
Garten Basel Bull. Vereins Freunde Zoo/. Gartens
Basel 43:\\-\3.
Nelson, E.W. 1876. additions to the avi-fauna of
Illinois with notes on other species of Illinois
birds. Bull. Nuttall Orn. Club 1:39-44.
Nesbitt, S.A. 1973. Wood Stork nesting in north
Florida. Fla. Field Nat. 1:25-26.
Nesbitt, S.A., and J.H. True. 1974. A new north
Florida Wood Stork colony. Fla. Field Nat.
2:14-15.
Nesbitt, S., W. Hetrick, and L. Williams, Jr.
1974. Foods of White Ibis from seven
collection sites in Florida. Proc. Annu. Conf.
Southeast. Assoc. Game Fish Comm. 28:517-532.
Nesbitt, S.A., J.C. Ogden, H.W. Kale, II, B.W.
Patty, and L.A. Rowse. 1982. Florida atlas of
breeding sites for herons and their allies: 1976-1978.
U.S. Fish Wildl, Serv., Office of Biol. Services,
FWS/OBS-81/49. 450pp, Washington DC.
Netchaev, V.A. 1963. [New data about the birds
of Low stream of Amur River.] Ornitologiya
6:177.
Neto, P.S. 1982. Aspectos bionomicos e
desenvolvimento de Theristicus caudatus
(Boddart 1783) (Aves, Threskiornithidae).
Dusenia 13(4): 145-149.
Neufeldt, LA., and K. Wunderlich. 1978.
Nipponia nippon (Temminck). In Atlas der
Verbreitung Palaearktischer Voegel, 7. Lieferung.
Akademie-Verlag, Berlin.
Neufeldt, LA., and K. Wunderlich. 1982. Ciconia
boyciana Swinhoe. In Stresemann, E., and L. A.
Portenko (eds), Atlas der Verbreitung
Palaearktischer Voegel. Akademie-Verlag, Berlin.
Neumann, O. 1928. Neue Formen von Nordost-
und Ost-Afrika. J. Orn. 76:783-787.
Neve de Mevergnies, G. 1988. Consommation de
Gerris et d'autres insectes par une cigogne
noire (Ciconia nigra). Aves 25:190-191.
Newman, K. 1974. Letter to the editor.
Honeyguide 79:48.
Newman, K. 1980. Birds of Southern Africa 1:
Kruger National Park. MacMillan,
Johannesburg.
361
Newman, K., and M. English. 1975. Openbill
Storks breeding in the Transvaal. Ostrich
46:264.
Newton, A. 1896. A Dictionary of Birds. A.&C.
Black, London. 1088pp.
Nice, M.M. 1933. The theory of territorialism
and its development. Pp. 89-100 in Fifty Years3
Progress of American Ornithology 1883-1933.
American Ornithologist's Union, Lancaster,
Pennsylvania.
Nice, M.M. 1943. Studies in the life history of the
Song Sparrow. II. The behavior of the Song
Sparrow and other Passerines. Trans. Linn. Soc
(AT) 6:1-329.
Nicholson, C.P. 1990. Recent records of Wood
Storks in Knox County, Tennessee. Migrant
61:10-11.
Nicholson, DJ. 1912. An Orange County Wood
Ibis rookery. Oologist 29:377-380.
Nicholson, DJ. 1929. Notes on the Roseate
Spoonbill (Ajaia ajaja) in Florida. Auk 46:381-
382.
Nicholson, E.M., I J. Ferguson-Lees, and P.A.D.
Hollom. 1957. The Camargue and the Goto
Donana. British Birds 50:497-519.
Nicholson, W.H. 1933. Roseate Spoonbills and
White Pelicans in Brevard County, Florida.
Auk 50:429.
Nicoll, M J. 1906. Sacred ibis from Dassen
Island, South Africa. Avic. Mag. 4(2):275-276.
Nicoll, M J. 1919. Handlist of the Birds of Egypt.
Ministry of Pub. Works, Egypt, Zool. Ser.,
Pub. 29, 120pp, Cairo.
Nieboer, E. 1965. Canthaxantin for Scarlet Ibises
Eudocimus ruber at Amsterdam Zoo. Int. Zoo Yb.
5:164-165.
Nielsen, B.P. 1969. Further spring observations
on the birds of Gilan, Northern Iran. Dan. Orn.
Foren. Tidsskr. 63:50-73.
Nielsen, B.P., and H.-J. Speyer. 1967. Some
observations of birds in northern Iran. Dan.
Orn. Foren. Tidsskr. 61:30-39.
Niethammer, G. 1938. Handbuech der Deutschen
Vogelkunde, vol. 2. Akademische
Verlagsgesellschaft, Leipzig.
Niethammer, G. 1972a. Storche ueber
Afghanistan. Z. Kolner Zoo 15:47-54.
Niethammer, J. 1972b. Ruckmeldung beringter
Turkestan-WeiBstorche (Ciconia c. asiatica) aus
Afghanistan. Vogelwarte 26:355.
Niethammer, G. 1967a. Storche in Afghanistan.
Vogelwarte 24:42-44.
Niethammer, J. 1967b. Zweijahre
Vogelbeobachtungen an stehenden Gewassern
bei Kabul in Afghanistan./ Orn. 108:119-
164.
Nikolaus, G. 1984. Distinct status changes of
certain Palearctic migrants in the Sudan.
Scopus 836-838.
Nikolaus, G. 1987. Distribution atlas of Sudan
birds with notes on habitat and status. Bonn.
Zool. Monogr. 25.
Nikolaus, G. 1989. Die Situation des Weiss-
Storchs im Sudan. Pp. 337-340 in Rheinwald,
G., J. Ogden, and H. Schulz (eds), Weiss-
Storch: Status und Schutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Nisbet, I.C.T. 1968. The utilization of
mangroves by Malayan birds. Ibis 110:348-
352.
Nisbet, I.C.T., and T.C. Smout. 1957. Autumn
observations on the Bosphorus and
Dardanelles. Ibis 99:483-499.
Nisbet, I.C.T., P.R. Evans, and P.P. Feeny.
1961. Migration from Morocco into southwest
Spain in relation to weather. Ibis 103a:349—
372.
Nitzsch, C.L. 1840. System der Pterylographie.
Eduard Anton, Halle.
Niven, C.K., and J.P.M. Niven. 1966a. Openbill
Stork Anastomus lamelligerus. Ostrich 37:58.
Niven, O.K., and J.P.M. Niven. 1966. Saddlebill
Stork feeding young. Ostrich 37:58-59.
Nixon, A. 1988. A further breeding record of the
White Stork in the southern hemisphere? Bee-
eater 39(4):55-56.
Noll, F.C. 1876. Besuch einiger Zoologischen
Garten und Aquarien im Juli 1876. ZooL Gart.
17:335-339.
Nores, M., and D. Yzurieta. 1980. Aves de
Ambientes Acudticos de Cordobay Centro de
Argentina. Academia Nacional de Giencias de
Cordoba, Argentina.
Norris, R.A. 1963. Birds of the A.E.C. Savannah
River Plant area. Contributions from the
Charleston Museum XIV, Charleston, USA.
Norris, R.A., and M.N. Hopkins, Jr. 1953.
Migration of birds in Ben Hill, Tift, and Irwin
Counties, Georgia. Oriole 18:1-9.
North, AJ. 1913-1914. Nests and Eggs of Birds
Found Breeding in Australia and Tasmania, vol. 4.
Trustees Austr. Mus., Sydney. 472pp.
North, M.E.W. 1939. Field notes on certain
raptorials and waterbirds in Kenya Colony.
Ibis (ser. 14) 3:487-507.
North, M.E.W. 1940. Field notes on Abdim's
Stork in two Kenya provinces, y. East Afr.
Uganda Nat. Hist. Soc. 15:1-5.
North, M.E.W. 1943. The breeding of the
Marabou Stork in East Africa. Ibis 85:190-
198.
North, M.E.W., and D.S. McChesney. 1964.
More Voices of African Birds. LP 33-1/3 rpm
record. Houghton Mifflin, Boston.
Norwood, J.R. 1956. Glossy Ibises in the
Mattamuskeet area. Chat 20:57-58.
Norwood, J.R. 1958. Wood Ibises inland in
North Carolina. Chat 22:44.
Noskiewicz, J. 1966. Results of inventory of
White Stork's nests in province Szczecin.
NotatkiOrn. 7:11-22.
Novaes, F.C. 1974. Ornitologia do Territorio do
Amapa, volume 1. Publicacoes Avulsas do
Museu Goeldi 25. Museu Paraense Emilio
Goeldi, Belem.
Nuttall, T. 1834. A Manual of the Ornithology of the
United States and of Canada. The Water Birds.
Hilliard, Gray, & Company, Boston.
Nutting, C.C. 1882. On a collection of birds from
the Hacienda 'La Palma', Gulf of Nicoya,
Costa Rica. Proc. US Natl. Mus. 5:382-409.
O'Reilly, J. 1958. The ordeal of the Wood Ibis.
Sports Illustrated SI March:51-54.
Oates, E.W. 1883. Birds of British Burmah. Porter,
London.
Oatiey, T.B., 1989. Information from White
Stork ring recoveries in Southern Africa. Pp.
357-360 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985. Schriftenr. DDA, Braunschweig,
Germany.
Oatiey, T.B., and M.A.M. Rammesmayer. 1988.
Review of recoveries of ringed White Storks
Ciconia ciconia in southern Africa. Ostrich 59:97-
104.
Ober, F.A. 1874. Birds of Lake Okeechobee.
Forest and Stream 2:162-163.
Oberholser, H.C. 1938. Bird Life of Louisiana.
Bull. Dep. Cons. State Louisiana 28.
Oberholser, H.C. 1945. Birds of the White-Fuller
expedition to Kenya, East Africa. Sci. Publ.
Cleveland Mus. Nat. Hist. 4(3):43-121.
Oberholser, H.C., and E.B. Kincaid, Jr. 1974.
The Bird Life of Texas. University of Texas
Press, Vol. 1, Austin, Texas, 530pp.
Odening, K. 1961. Zur Kenntnis der
Trematodenfauna einiger Vogelarten. Mitt.
ZooL Mus. Bed. 37:125-146.
Odom, R.R. 1976. 1975 heronry survey of the
Georgia coast. Oriole 41:19-35.
Odom, R.R. 1978. Wood Storks nesting on the
Georgia coast. Oriole 43(1): 1-5.
Odom, R.R., J.W. Guthrie, J.H. Rappole, and
B J. Freeman. 1979. Wood Stork breeding
colony in Glynn County, Georgia. Oriole
44:88-89.
Officer, R.H. 1958. Birds of the R.A.O.U. 1957
Camp-out Portland District, Victoria. Emu
58:375-390.
Ogasawara, K., and Y. Izumi. 1977. Japanese
White Storks (Ciconia ciconia boyciana) appeared
in northern Honshu and Hokkaido. Misc. Rep.
Yamashina Inst. Orn. 9:121-127.
Ogden, J.C. 1976. The pink pause, a previously
undescribed behavior by Roseate Spoonbills.
Fla. Field Nat. 4:34-35.
Ogden, J.C. 1978a. Recent population trends of
colonial wading birds on the Atlantic and Gulf
coastal plains. Pp. 137-153 in Sprunt, A., J.C.
Ogden, and S. Winckler (eds), Wading Birds.
Research Report No. 7, National Audubon
Society, New York.
Ogden, J.C. 1978b. Endangered Wood Stork. Pp.
3-4 in Kale, H.W. (ed.), Rare and Endangered
Biota of Florida, vol. 2, Birds. University Presses
of Florida, Gainesville.
Ogden, J.C. 1978c. Roseate Spoonbill. Pp. 52-54
in Kale, H.W. (ed.) Rare and Endangered Biota of
Florida, vol. 2, Birds. University Presses of
Florida, Gainesville.
Ogden, J.C. 1981. Nesting distribution and
migration of Glossy Ibis Plegadis falcinellus in
Florida, U.S.A. Fla. Field Nat. 9:1-6.
Ogden, J.C. 1985. The Wood Stork. Pp. 458-470
in Di Silvestro, R.L. (ed.), Audubon Wildlife
Report 1985. National Audubon Society, New
York.
Ogden, J.C., and S.A. Nesbitt. 1979. Recent
Wood Stork population trends in the United
States. Wilson Bull. 91:512-523.
Ogden, J.C., and B. Patty. 1982. The recent
status of the Wood Stork in Florida and
Georgia. Pp. 97-103 in R. Odom and J.
Gutherie (eds.), Proceedings of the Nongame and
Endangered Wildlife Symposium. Georgia Dept
Natural Resource Technical Bulletin WL5.
Ogden, J.C., and B.T. Thomas. 1985a. An
opened wing foraging behavior by the Green
Ibis. Colonial Waterbirds 8:181-182.
Ogden, J.C., and B.T. Thomas. 1985b. A
colonial wading bird survey in the central
llanos of Venezuela. Colonial Waterbirds 8:23-
31.
Ogden, J.C., J.A. Kushlan, and J.T. Tilmant.
1976. Prey selectivity by the Wood Stork.
Condor 78:324-330.
Ogden, J.C., J. A. Kushlan, and J.T. Tilmant.
1978. The food habits and nesting success of
wood storks in Everglades National Park,
1974. U.S. Natl. Park Serv. Nat. Resour. Publ. 16.
Ogden, J.C., J.A. Rodgers, R. Andrews, S.A.
Nesbitt, M.C. Landin, and J.A. Kushlan.
1980. Current management of colonial
waterbirds in the United States. Pp. 32-38 in
Parnell, J. and R.F. Soots (eds), The
Management of Colonial Waterbirds. North
Carolina Sea Grant Publication UNC-SG-80-
06, Wilmington.
Ogden, J.C., D.A. McCrimmon, Jr., G.T.
Bancroft, and B.W. Patty. 1987. Breeding
populations of the Wood Stork in the
southeastern United States. Condor 89:752-
759.
Ogilvie-Grant, W.R. 1889. On the genus Platalea,
with a description of a new species from New
Guinea. Ibis (ser. 6) 1:32-58.
Ogilvie-Grant, W.R. 1902. A new species of
stork, of the genus Dissura. Bull. Br. Orn. Club
13:26-27.
Ogilvie-Grant, W.R. 1903. On a new stork from
Borneo. Ibis (ser. 8) 3:145-148.
Ogilvie-Grant, W.R., and J.D.D. La Touche.
1907. On the birds of the island of Formosa.
Part II. Ibis (ser. 9) 2:254-279.
362
Ogle, D. 1986. The status and seasonality of
birds in Nakhon Swan Province, Thailand.
Nat. Hist. Bull. Siam Soc. 34:115-143.
Ohata, K. 1991. Additional records of the
Oriental White Storks from Lake Utonaito,
Japan. Spec. Group Storks, Ibises, Spoonbills
4(2):3-4.
Ohlendorf, H.M., E.E. Klaas, and T.E. Kaiser.
1978. Organochlorine residues and eggshell
thinning in Wood Storks and Anhingas. Wilson
Bull. 90:608-618.
Ohlendorf, H.M., E.E. Klaas, and T.E. Kaiser.
1979. Environmental pollutants and eggshell
thickness: Anhingas and wading birds in the eastern
United States. U.S. Fish Wildl. Serv. Spec. Sci.
Rep. Wildl. 216, Washington DC.
Oleinikov, N.S., B.A. Kazakov, N.K. Lomandze,
and I.M. Yaxkova. 1975. The ibis family in
Ciscaucasia USSR. Izv. Sev., Kavk. Nauchn.
Tsentra. Vyssh. Shk. Estestv. Nauki. 3:52-54.
Olivares, A. 1969. Aves de Cundihamarca. Univ.
Nacional de Columbia, Direction de
Divulgacion Cultural, Bogota.
Oliveira Pinto, O.M. de. 1964. Ornitologia
brasiliense. Catalogo descritivo e ilustrado das aves do
Brasil, vol. I. Departamento de Zoologia da
secretaria de agricultura do Estado de Sao
Paulo, Sao Paulo.
Oliver, J.D., and S.A. Schoenberg. 1989.
Residual influence of macronutrient
enrichment on the aquatic food web of an
Okeefenokee Swamp abandoned bird rookery.
(M<w 55:175-182.
Oliver, W.L.R., M.M. Mallet, D.R. Singleton,
and J.S. Ellett III. 1979. Observations on the
reproductive behaviour of a captive colony of
Bare-faced Ibis Geronticus eremita. Dodo 16:11-
35.
Oliver, W.R.B. 1955. New Zealand Birds.
A.H.&A.W. Reed, Wellington.
Olmos, F. 1990. Nest predation of Plumbeous
Ibis by Capuchin Monkeys and Black Hawks.
Wilson Bull. 102:169-170.
Olney, P. 1986. London Zoo-1985. Avic. Mag.
92:110-111.
Olrog, C.C. 1959. Las aves Argentinas. Una guia de
campo. Inst. Miguel Lillo, Tucuman. 343 pp.
Olrog, C.C. 1963. Listay Distribucion de las Aves
Argentinas. Op. Lilloana 9. Tucuman,
Argentina. 377 pp.
Olrog, C.C. 1965. Diferencias en el ciclo sexual
de algunas aves. Hornero 10:269—272.
Olrog, C.C. 1968. Las Aves Sudamericanas, vol. 1.
Universidad National, Tucuman.
Olrog, C.C. 1975. Vagrancy of Neotropical
cormorants, egrets, and White-faced Ibis. Bird
Banding 46:207-212.
Olson, S.L. 1974. Tantalus milneedwardsii
Shufeldt—a synonym of the Miocene pheasant
Miophasianus altus (Milne-Edwards). Wilson
Bull. 86:110-113.
Olson, S.L. 1979. Multiple origins of the
Ciconiiformes. Proc. 1978 Conf. Colonial
Waterbird Group 2:165-170.
Olson, S.L. 1981. The generic allocation of Ibis
pagana Milne-Edwards, with a review of fossil
ibises (Aves: Threskiornithidae).y. Vertebr.
Paleontol. 1:165-170.
Olson, S.L. 1985a. Early Pliocene ibises (Aves,
Plataleidae) from south-western Cape
Province, South Africa. Ann. S. Afr. Mus.
97:57-69.
Olson, S.L. 1985b. The fossil record of birds. Pp.
79-238 in Farner, D.S., J.R. King, and K.C.
Parkes (eds), Avian Biology, vol. 8. Academic
Press, New York.
Olson, S.L., and A. Feduccia. 1980.
Relationships and evolution of flamingos
(Aves: Phoenicopteridae). Smithson Contr. ZooL
316: 1-73.
Olson, S.L., and H.F.James. 1984. The role of
Polynesians in the extinction of the avifauna of
 the Hawaiian Islands. Pp. 768-780 in Martin,
P.S., and R.G. Klein (eds), Quartenary
Extinctions: a Prehistoric Revolution. University of
Arizona Press, Tucson.
Olson, S.L. and H.F. James. 1991. Descriptions of
Thirty-two new species of birds from the Hawaiian
Islands. Part I. Non-passeriformes. Ornithological
Monographs 45:1-88.
Olson, S.L., and D.W. Steadman. 1977. A new
genus of flightless ibis (Threskiornithidae) and
other fossil birds from cave deposits in
Jamaica. Proc. Bioi Soc. Wash. 90:447-457.
Olson, S.L., and D.W. Steadman. 1979. The
humerus ofXenicibis, the extinct flightless ibis
ofJamaica. Proc. Biol. Soc. Wash. 92:23-27.
Olsen, S.L., and A. Wetmore. 1976. Preliminary
diagnoses of two extraordinary new genera of
birds from Pleistocene deposits in the
Hawaiian Islands. Proc. Biol. Soc. Wash.
89:247-258.
Olson, S., M. Rea, and P. Brodkorb. 1986.
Comment on the proposed grant of precedence
to Threskiornithidae Richmond, 1917 (Aves)
over Plataleinae Bonapart, 1938.
Z.N.(S.)2136. Bull. Zoo/. Norn. 43:10-11.
Olsson, J., R. Asterling, and L. Larsson. 1980.
White-tailed black storks. British Birds 73:104.
Omaston, A.E. 1913. The birds of Gorakhpur.J.
Bombay Nat. Hist. Soc. 22:532-549.
D'Ombrain, E.A. 1906. Visit to an ibis rookery.
Emu 5:185-189.
Oordt, G.J. v. 1937. Die dritte hollandische
Loefflerkolonie. Beitr. Fortpfanzungsbiol. Vogel
13.
Oordt, G.J. v. 1939. De Lepelaarkolonie in de
Muy.Ardea 28:113-114.
Oordt, G.J. v. 1954. The Eider Duck and the
Spoonbill in the Netherlands. Oryx 2:379-380.
Oordt, G.J. v. 1957. The Spoonbill in the
Netherlands. Pp. 30-35 in Bannerman, D.A.
(ed.), The Birds of the British Isles, vol. 6. Oliver
and Boyd, Edinburgh.
Ord, G. 1817. An account of an American
species of the genus Tantalus or Ibis. Acad. Nat.
Set. Phila. 1:53-57.
Oreel, G.J. 1980. De Zwarte Ooievaar. Lepelaar
68:104-105.
Ornithological Society of Japan. 1975. Checklist of
Japanese Birds, 5th edn. Gakken, Tokyo. 364pp.
Ornithological Sub-Committee of the East
African Natural History Society. 1977. Some
recent records of Palaearctic birds in Kenya
and Tanzania. Scopus 1:39-43.
Ornithological Sub-Committee of the East
African Natural History Society. 1978. East
African bird report 1977. Scopus 1:114-131.
Ornithological Sub-Committee of the East
African Natural History Society. 1979. East
African bird report 1978. Scopus 2:105-144.
Ortiz Crespo, F., and S. Valarezo Delgado. 1975.
Lista de aves del Ecuador. Publicaciones de la
Sociedad Ecuatoriana Francisco Campos 2,
Guayaquil.
Osborn, R.G., and T.W. Custer. 1978. Herons and
Their Allies: Atlas of Atlantic Coast Colonies, 1975
and 1976. U.S. Fish Wildl. Serv. Biol. Rep.
FWS/OBS 77/08. U.S. Fish and Wildlife
Service, Laurel, Maryland.
Osieck, E.R., and C.N. deVries, 1987. Lepelaar
vooruit. [Increase of Spoonbills.] Limosa
60:155.
Ossowski, L.L.J. 1952. The Hadedah Ibis,
Hagedashia hagedash hagedash (Latham), and its
relation to pest control in wattle plantations.
Ann. Natal Mus. 12:279-290.
Ostermann, W. 1987 [Breeding record of Ciconia
nigra in eastern Westphalia.] Charadrius 23:208.
Ott, J., and P. Joslin. 1981. A helping hand back
to mother. Brookfield Bison Feb/Mar:Cl-C8.
Oustalet, E. 1872. Remarques sur 1'ibis sinensis.
Paris Mus. Nat. d'Hist. Nat. Archiv. 8:129-137.
Oustalet, M. 1865. Observations sur le groupe
des ibis. Paris Mus. Nat. d' Hist. Nat. Archiv.
1(2).
Owen, T.R.H. 1958. The Shoebill Stork. Afr.
Wild Life 12(3):\9\-195.
Paakspuu, V. 1977. [Observations of storks and
common herons at Matsalu Bay in summer,
1976.] Loodusvaatlusi (1976) 1:125-127.
Packard, F.M. 1947. Notes on the occurrence of
birds in the Gulf of Mexico. Auk 64:130-131.
Paget-Wilkes, A.H. 1928. The birds of the region
south of Lake Nyasa. Part 1. Ibis (ser. 12)
4:690-748.
Paige, J. P. 1960. Bird notes from Aden and
Oman. Ibis 102:520-525.
Paine, J. 1989. The Yok Don Reserve (Daklak).
Garrulax 6:9-11.
Pala, S. 1971. Faydal: kuslarimizdan
(Geronticus— Comatibis — eremita L.) — ozellikleri,
yasayisi ve beslenmesi. Tabiat ve Insan 5:14—22.
Pallary, P. 1922. Note sur le regime alimentaire
de 1'ibis a tete chauve (Geronticus eremita L.), un
oiseau marocain a proteger. Bull. Soc. Set. Nat.
maroc2(12):48.
Palmer, R.S. 1962. Handbook of North American
birds, Volume 1, Loons through Flamingos. Yale
University Press, New Haven, Connecticut.
Palmisano, A.W. 1971. Ibis in Louisiana.
Louisiana Wildl. Fish. Comm. Wildl. Tech. Rep.
71:1-22.
Paludan, K. 1959. On the birds of Afghanistan.
Vidensk. Medd. Dan. Naturhist. Foren. 122.
Panday, J.D. 1974. Storks preying on live birds.
J. Bombay Nat. Hist. Soc. 71:141.
Pankin, N.S., and LA. Neufeldt. 1976. The Far
Eastern White Stork in Amur Region. Trans.
Oka State Nat. Res. 12:19-31.
Panouse,J.B. 1951. Sur les migrations des
Cigognes Blanches d' Europe occidentale et du
Maroc. C.R. Seances Acad. Set. Paris 232:557-
559.
Parasharya, B.M., and R.M. Naik. 1988.
Ciconiiform birds breeding in Bhavnagar City,
Gujarat. J. Bombay Nat. Hist. Soc. Centenary
Symp.
Parent, G.H. 1973. La signification ecologique de
la nidification de la cigogne blanche (Ciconia
ciconia L.) en Lorraine, Beige, en 1972. Aves
10:70-112.
Parish, D. 1984. Stork Report. Interwader 1983
Report. Interwader Publications, Kuala
Lumpur, Malaysia.
Parish, D. 1985. Ground surveys in peninsular
Malaysia. Interwader 1984 Report, pp. 64-78.
Interwader Publications, Kuala Lumpur,
Malaysia.
Parish, D., and D.R. Wells (eds). 1984. Interwader
1983 Report. Interwader Publications, Kuala
Lumpur, Malaysia.
Parish, D., R.C. Prentice, and C.E. Taylor. 1986.
Foraging behaviour of the Milky Stork.
Interwader Publ. 16:27, Kuala Lumpur,
Malaysia.
Parker, I.S.C. 1982. Two observations of nesting
in the eastern race of the Green Ibis Bostrychia
olivacea akeleyorum. Scopus 6:20.
Parker, I.S.C. 1984a. Shoebill Balaeniceps rex: A
deletion from the Kenya avifauna. Scopus 8:79.
Parker, I.S.C. 1984b. Shoebill Balaeniceps rex and
Wattled Crane Grus carunculatus in the
Moyowosi Swamp, Tanzania. Scopus 8:24—25.
Parker, S.A., et al. 1979. An annotated checklist of
the birds of South Australia. Part 1: Emus to
Spoonbills. S. Aust. Orn. Assoc., Adelaide.
Parker, T.A., and S.A. Parker. 1982. An Annotated
Checklist of Peruvian Birds. Buteo Books,
Vermillion, South Dakota.
Parker, W.K. 1860. Abstract of notes on the
osteology of Balaeniceps rex. Proc. Zool. Soc. Lond.
1860:324-330.
Parker, W.K. 1864. On the sternal apparatus of
birds and other vertebrata. Proc. Zool. Soc.
Lond. 1864:339-341.
363
Parker, W. 1985. Creating Wood Stork habitat:
an important management strategy. Endang.
Species Tech. Bull. 10:7.
Parkes, K.C. 1951. The generic name of the
White and Scarlet Ibises. Proc. Biol. Soc. Wash.
64:61.
Parkes, K.C. 1955. Systematic notes on North
American birds.I. The herons and ibises
(Ciconiiformes). Ann. Carnegie Mus. 33:287-
293.
Parkes, K.C. 1978. A review of the classification
of Ciconiiformes. Pp. 7-15 in A. Sprunt, IV,
J.C. Ogden, and S. Winckler (eds), Wading
Birds. Research Report No. 7, National
Audubon Society, New York.
Parmalee, P.W., and G. Perino. 1970. A
prehistoric archeological record of the Roseate
Spoonbill in Illinois. Trans. III. State Acad. Sci,
63:254-258.
Parnell, F.I. 1942. The Bald Ibis in Basutoland.
Oj/wA 13:100-101.
Parnell, J.F., and R.F. Soots, Jr. 1979. Atlas of
colonial waterbirds of North Carolina estuaries.
UNC Sea Grant, Publication UNC-SG-78-10.
Parnell, J.F., D.G. Ainley, H. Blokpoel, B. Cain,
T.W. Custer, J.L. Dusi, S. Kress, J.A.
Kushlan, W.E. Southern, L.E. Stenzel, and
B.C. Thompson. 1988. Colonial waterbird
management in North America. Colonial
Waterbirds \ 1:129-169.
Parslow, J.L.F. 1973. Organochlorine insecticide
residues and food remains in a Bald Ibis
Geronticus eremita chick from Birecik, Turkey.
Bull. Br. Orn. Club 93:163-166.
Parsons, J. 1976. Counts of ciconiiform nesting in
three colonies near Kisumu heronry—27 April
1976. Bull. E. Afr. Nat. Hist. Soc. 1976:61-62.
Parsons, J. 1977. The effect of predation by Fish
Eagles on the breeding success of various
Ciconiiformes nesting near Kisumu, Kenya. J.
Nat. 7/^.11:337-353.
Parsons, P.F. 1974. Openbill Storks nesting.
Honey guide 78:41-43.
Passburg, R.E. 1959. Bird notes from northern
Iran. 7£w 101:153-169.
Patnaik, M.M., A.T. Rao, L.N. Acharjyo, and
D.N. Mohanty. 1970. Notes on a nodular
disease of the intestine of the Open-billed
Stork (Anastomus oscitans) caused by
Chaunocephalusferox.J. Wildl. Dis. 6:64-66.
Patterson, T.K. 1977. Population of an egret
roost in East Dublin, Georgia. Oriole 42:26-29.
Patterson, T.K. 1979. Heronry located in
Laurens County, Georgia. Oriole 44:55—56.
Paulian, R. 1955. Les animaux proteges de
Madagascar. Tananarive, Tsimbazaza.
Pauraj, S., and G. Gunasekaran. 1988. The
Vedanthangal Water-bird Sanctuary: a new
breeding ground for Pelicans and Painted
Storks. J. Bombay Nat. Hist. Soc. 85:414-^15.
Paynter, R.A. 1955. The Ornithogeography of the
Yucatan Peninsula. Peabody Mus. Nat. Hist.,
Yale Univ. Bull. 9, New Haven.
Paynter, R.A., Jr., M.A. Taylor, Jr., and B.
Winter. 1975. Ornithological Gazetter of Bolivia.
Harvard Press, Cambridge, Massachusetts.
Paz, U. 1984. [Even the Stork in heavens knows
her times.] Sunbird2:\\-\6.
Paz, U. 1987. The Birds of Israel. C. Helm,
London.
Pearson, D.L., and M.A. Plenge. 1974. Puna
bird species on the coast of Peru. Auk 91:626-
631.
Pearson, D.J., and D.A. Turner. 1986. The less
common Palaearctic migrant birds of Uganda.
Scopus 10:61-82.
Pearson, T.G. 1892. The ibises of Ledworth
Lake, Florida. Oologist 9:99-100.
Pearson, T.G. 1911. Report of the Secretary. 'Bird
Lore 14:383-392.
Pearson, T.G. 1919. Roseate Spoonbill in North
Carolina. Auk 36:568.
Pearson, T.G. 1921. Notes on the bird-life of
southeastern Texas. Auk 38:513-523.
Pearson, T.G. 1922. Notes on the birds of
Cumberland Island, Georgia. Wilson Bull.
34:84-90.
Pearson, T.G. 1925. White Ibis. Bird Lore 27:75-
78.
Pearson, T.G. 1936. Birds of America. Garden City
Publ. Co., New York.
Pedrero, P.C. 1983. Estudio general sobre la
dinamica de la poblacion de la Ciguena
Blanca, Ciconia c. ciconia (L.) en Espana. Thesis
Doctoral, Universidad Complutense de
Madrid.
Peet, T.E., and W.L.S. Loat. 1913. The ibis
cemetery. Pp. 40-47 in The Cemeteries ofAbydos,
vol. 3. Egypt Exploration Society, London.
Pegoraro, K. 1985. Behavioral development of
young Hermit ibis females. J. Orn. 126:19-23.
Pegoraro, K. 1986. Hat der Waldrapp chancen,
seine Wiederausburgerung zu erleben [abstr.].
Deutchen Ornithologen-Gesellschaft 27 September
1986.
Pegoraro, K. 1989. Waldrapp Ibis. Spec. Group
Storks, Ibis Spoonbills 2:10-11.
Pegoraro, K. and G. Malin. 1990. Bald Ibis
(Geronticus eremita) in Morocco—attendance of
immature birds at breeding colonies. J. Orn.
131:453-456.
Pegoraro, K., and E. Thaler. 1985. Zum
Verhalten erstbrutender Waldrapp-Weibchen
im Alpenzoo. Zool. Gart. 55(2/3): 113-123.
Pegoraro, K., and E. Thaler. 1990. Stop the
extinction of the Waldrapp ibis! Pp. 173-174
in Dresser, B.L. et al. (eds), Proceedings 5th
World Conference on Breeding Endangered Species in
Captivity, Cincinnati, Ohio.
Peirce, M.A., and J.E. Cooper. 1977.
Haematozoa of East African birds. IV.
Redescription of (Haemoproteus crumenium), a
parasite of the Marabou Stork. E. Afr. Wildl.J.
15(2): 169-172.
Pelle, J. 1989. Status und Verbreitung des Weiss-
Storchs injugoslawien 1984. Pp. 115-120 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Pelzeln, A. v. 1868-1871. Zur Ornithologie
Brasiliens. Pichlers Witwe and Sohn, Wein.
Pelzer, J. 1972. The fabled storks they range the
world. Int. Wildl. 2:42-44.
Penard, P.P., and A.P. Penard. 1908. De Vogels
van Guyana, vol. I. P.P. Penard, Paramaribo,
Guyana.
Penard, T.E. 1926. Scarlet Ibis. In Bent, A.C.
Life histories of North American marsh birds.
USNatl. Mus. Bull. 135:33-45.
Pence, D.B. 1971. The hypopi (Acarina:
Hypoderidae) from the subcutaneous tissues of
the White Ibis, Eudocimus albus L. J. Parasitol.
57:1321-1323.
Pence, D.B. 1972. The hypopi (Acarina:
Sarcoptiformes: Hypoderidae) from the
subcutaneous tissues of birds in Louisiana. J.
Med. Entomol. 9:435-438.
Pence, D.B., and A.O. Bush. 1973. Polycyclorchis
eudocimi gen. et sp. n. (Trematoda:
Cyclocoelidae) from the trachea of the White
Ibis (Eudocimus albus L.)./ Parasitol. 59:85-89.
Pennock, CJ. 1919. Notes on the birds of
Wakulla County, Florida. Wilson Bull. 31:109-
117.
Penny, M. 1974. The Birds of the Seychelles and the
Outlying Islands. Collins, London. 160pp.
Pennycuick, C J. 1969. The mechanics of bird
migration. Ibis 111:525-556.
Pennycuick, C J. 1972. Soaring behaviour and
performance of some East African birds,
observed from a motor-glider. Ibis 114:178-
218.
Pennycuick, C.J., and T. De Santo. 1989. Flight
speeds and energy requirements for White
Ibises on foraging flights. Auk 106:141-144.
Perdeck, A.C., and BJ. Speek. 1964. Ringverslag
van het vogeltrekstation. Limosa 37:96-186.
Perdeck, A.C., and BJ. Speek. 1965. Ringverslag
van het vogeltrekstation. Limosa 38:142—199.
Perdeck, A.C., and B J. Speek. 1966. Ringverslag
van het vogeltrekstation. Limosa 39:85-134.
Perdeck, A.C., and B J. Speek. 1967. Ringverslag
van het vogeltrekstation. Limosa 40:59-112.
Perennou, C. 1987. Two important wetlands
near Pondicherry. Blackbuck 3:3-11.
Perez Vigueraz, I. 1960. Notas sobre algunos
cestodes encontrados en Cuba. Libro
Homenaje al Dr. Eduardo Caballero y
Caballero.>«to> 1930-1960:377-397.
Pernetty, A.J. 1763. Histoire d'un Voyage aux Isles
Malouines. Assemblela Legislativa do Estado de
Santa Catarina—Assessoria Cultural, Saillant-
Delacroix and Lyon, Paris.
Perrault, C. 1734a. Description anatomique de
quatre pallettes (Platalea). Mem. Acad. Set. Paris
(1666-99), 3(3):22-30.
Perrault, C. 1734b. Description anatomique d'un
ibis blanc et de deux cicognes. Mem. Acad. Sci.
Paris (1666-99), 3(3):58-72.
Perrine, D. 1989. The world was never made for
someone as beautiful as you. Living Bird Q.
8(2):32-34.
Pescott, T. 1968. Ibis drown off Point Henry?
Australian Bird Watcher 3:130-132.
Peters, D.S. 1983. Die 'Schnepfenralle'
Rhynchaeites messelensis Wittich 1898 is ein Ibis.
[Rhynchaeites messelensis is an ibis.] J. Orn.
124:1-27.
Peters, H.B. 1928. Aus der Biologic des
Waldrapps. Naturforcher 5:380-383.
Peters, H. 1990. Waldrappd'dmmerung am Euphrat.
Kasparek Verlag, Heidelberg.
Peters, J.L. 1931. Check-list of Birds of the World.
Harvard University Press, Cambridge,
Massachusetts.
Peters, W. 1867. Ueber den Kehlsack des
Marabustorches (Leptoptilos crumeniferus). Mber.
Kon. Preuss. Akad. Wiss. Berlin 1866:168-169.
Peterson, P.C., and W.T. Atyeo. 1977. a new
genus and species of alloptine feather mite
from the Roseate Spoonbill (Acarina,
Analgoidea, Alloptidae). Steenstrupia
4(17): 189-193.
Peterson, R.T. 1934. A Field Guide to the Birds.
Houghton Mifflin Company, Boston.
Peterson, R.T. 1962. White Storks, vanishing
sentinels of the rooftops. Natl. Geogr. Mag.
121:838-853.
Peterson, R.T., and E.L. Chalif. 1973. A Field
Guide to Mexican Birds (and Adjacent Central
America). Houghton Mifflin Company, Boston.
Peterson, R.T., G. Mountfort, and P.A.D.
Hollom. 1983. A Field Guide to the Birds of
Britain and Europe. Houghton Mifflin Company,
Boston.
Peterson, T. 1976. Vit stork (Ciconia ciconia) och
svart stork (Ciconia nigra} pa. Oland. [The
White Stork (Ciconia ciconia) and the Black
Stork (Ciconia nigra) on the Oland.] Calidris.
76(3):79-84.
Petherick, J. 1860. Memoranda on the
Hippopotamus and Balaeniceps recently
imported to England, and now in the gardens
of the Zoological Society. Proc. Zool. Soc. Lond.
1860:195-199.
Petit, D.R., and K.L. Bildstein. 1986.
Development of formation flying in juvenile
White Ibis (Eudocimus albus). Auk 103:244-246.
Petit, D.R., and K.L. Bildstein. 1987. Effect of
group size and location within the group on
the foraging behavior of White Ibises. Condor
89:602-609.
Pettingill, O.S. Jr. 1971. A Guide to Bird Finding
East of the Mississippi. Houghton Mifflin
Company, Boston.
364
Petzold, H.G. 1958. Einige Bilder und Gedanken
zum Thema 'Kronismus beim WeiBstorch'.
Beitr. Vogelkde 6:261-265.
Phelps, F.M. 1912. A March bird-list from the
Caloosahatchee River and Lake Okeechobee.
Wilson Bull. 24:117-125.
Phelps, F.M. 1914. The resident bird life of the
Big Cypress Swamp region. Wilson Bull. 26:86-
101.
Phelps, W.H., and R. Meyer de Schauensee.
1979. Aves de Venezuela. Princeton University
Press, Princeton.
Phelps, W.H., and W.H. Phelps, Jr. 1958. Lista
de las aves de Venezuela y su distribucion.
Tomo II, Parte I. No Passeriformes. Sep. Bol.
Soc. Venezolana Ciencias Nat. 19(90): 1-317.
Philippona, J. 1967. Short notes. The Carska
Bara in Yugoslavia. Ardea 55:146-147.
Phillips, A.R. 1968. Geologic age of Ciconia
maltha. Auk 85:315.
Phillips, A.R. 1986. Comment on the proposed
grant of precedence to Threskiornithidae
Richmond, 1917 (Aves) over Plataleinae
Bonapart, 1838. Z.N.(S.)2136. Bull. Zool. Norn.
43:10-11.
Phillips, W.W.A. 1940. The occurrence of the
Black Stork (Ciconia nigra) in Ceylon. Ibis (ser.
14) 4:333-334.
Phillips, W.W.A. 1978. Annotated Check List of the
Birds of Ceylon (Sri Lanka). Wildlife and Nature
Protection Society and Ceylon Bird Club,
Colombo, Sri Lanka. 92pp.
Phillips, W.W.A., H. Rosenberg, E. Schuz, and
H. Seilkopf. 1959. Zum Zug des Weiss-Storchs
im Gebiet von Rotem Meer, Suesgolf und
Kanalzone. Vogelwarte 20:116-121.
Picken, V.B. 1987. The Hong Kong Bird Report
198411985. The Hong Kong Bird Watching
Society, Hong Kong. 127pp.
Piechocki, R. 1968. Beitrage zur Avifauna der
Mongolei. Teil 1. Non-Passeriformes. Mitt.
Zool. Mus. Berl. 44:149-292.
Piekarz, R. 1965. Breeding White Storks in
Warsaw Zoo. Int. Zoo Yb. 5:126-127.
Pierre, P. 1988. Status actuel de la Cigogne Noire
(Ciconia nigra) en Wallonie. Aves 25:183-189.
Piesker, O. 1967. Zum Horstbau-Verhalten von
Weiss-Stoerchen. Falke 14:206-207.
Pike, E.R. 1931. Roseate Spoonbills in Florida.
Auk 48:423-424.
Pineau, J., and M. Giraud-Audine. 1977. Notes
on nesting birds from the extreme northwest of
Morocco, reproduction and movements.
Alauda 45:75-103.
Pinowska, B., and J. Pinowski. 1989. Feeding
ecology and diet of the White Stork Ciconia
ciconia in Poland. Pp. 381-396 in Rheinwald,
G., J. Ogden, and H. Schulz (eds), Weiss-
Storch: Status undSchutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Pinowski, J., and G. Morales. 1981. Aspectos
ecologicos de las aves de los modulas de apure.
Bol. Soc. Venezolana Cienc. Nat. 35:67-78.
Pinowski, J., L.G. Morales, J. Pacheco, K.A.
Dobrowolski, and B. Pinowska. 1980.
Estimation of the food consumption of fish-
eating birds in the seasonally-flooded
savannahs (llanos) of Alto Apure, Venezuela.
Bull. Acad. Pol. Sci. Ser. Sci. Biol. 28:163-170.
Pinto, A.A. da Rosa. 1983. Ornitologia de Angola.
Institute de Investigacao Cientifica Tropical,
Lisboa.
Pinto, O.M. deO. 1964. Ornitologia Brasiliense,
vol. 1. Dept. da Zoologia da Secretaria da
Agricultura, Sao Paulo, Brazil.
Pitman, C.R.S. 1928. The nesting ofHagedashia
hagedash nilotica — the Nile Valley Hadada in
Uganda. Ool. Rec. 8:44-46.
Pitman, C.R.S. 193la. The nesting of Dissoura
episcopus microscelis (Sclater) the African
 Woolly-necked Stork in Uganda. OoL Record
11:45-46.
Pitman, C.R.S. 193Ib. Further notes on the
breeding of Hagedashia hagedash nilotica—the
Nile Valley Hadada in Uganda. OoL Rec.
11:48.
Pitman, C.R.S. 1945. Some breeding colonies of
the Marabou Stork in Uganda. Ibis 87:282-
283.
Pitman, C.R.S. 1957. Further notes on aquatic
predators of birds. Part II. Bull. Br. Orn. Club
77:105-110.
Pitman, C.R.S. 1961. Birds perching on
hippopotamus. Bull. Br. Orn. Club. 81:148.
Pitman, C.R.S. 1965. The nesting, eggs and
young of the Saddle-bill Stork, Ephippiorhynchus
senegalensis (Shaw). Bull. Br. Orn. Club 85:70-
80.
Pizzey, G. 1980. A Field Guide to the Birds of
Australia. Princeton University Press,
Princeton.
Plath, L. 1979. Loeffler (Platalea leucorodia L.) an
der Mittelelbe im Kreis Havelberg. [Spoonbill
(Platalea leucorodia) in the Middle Elbe at Kreis
Havelberg.] Beitr. Vogelkde 25(1/2): 124.
Plattner, F. 1956. Die Verbreitung des WeiB-
storches im Gebit des Urmiasees (Iran).
VogelwartelS:\78-\79.
Plowes, D.C.H. 1947. The birds of Ladysmith,
Natal. Ostrich 18:134-154.
Pocock, T.N., and C J. Uys. 1967. The Bald Ibis
in the north-eastern Orange Free State.
Bokmakierie 19:28-31.
Podmore, C.R. 1957. In the Sudan and Western
Kenya with J.G. Millais. Afr. Wildl. 11:235-
243.
Pollard, C J.W. 1981. A vulture restaurant at the
Victoria Falls. Honeyguide 107/108:39.
Pomeroy, D.E. 1973. The distribution and
abundance of Marabou Storks in Uganda. E.
Afr. Wildl. J. 11:227-240.
Pomeroy, D.E. 1975a. Birds as scavengers of
refuse in Uganda. Ibis 117:69-81.
Pomeroy, D.E. 1975b. Marking Marabou Storks.
Bird-Banding 46:343-344.
Pomeroy, D.E. 1977a. Marabous associated with
vulture parties in East Africa. Scopus 1:103-
106.
Pomeroy, D.E. 1977b. The biology of Marabou
Storks in Uganda. I. Some characteristics of
the species, and the population-structure.
Ardea 65:\-W.
Pomeroy, D.E. 1977c. Marabou Stork breeding
colonies in Uganda. J. E. Afr. Nat. Hist. Soc.
Natl.Mus.3l:\-\\.
Pomeroy, D.E. 1978a. Counts of Marabou Storks
(Leptoptilos crumeniferus) in relation to their
movements in eastern Africa. Scopus 2:92-96.
Pomeroy, D.E. 1978b. The biology of Marabou
Storks in Uganda. II. Breeding biology and
general review. Ardea 66:1-23.
Pomeroy, D.E. 1978c. Seasonality of Marabou
Storks Leptoptilos crumeniferus in Eastern Africa.
/Aw 120:313-321.
Pomeroy, D.E. 1979. Nesting of Marabou Storks
in the Makindu-Kiboko-Kibwezi area. E. Afr.
Nat. Hist. Soc. Bull. 1979:114-116.
Pomeroy, D.E. 1986. The Marabou Stork in
Kenya. Scopus W:\-9.
Pomeroy, D.E. 1989. The White Stork in Kenya.
Pp. 355-356 in Rheinwald, G., J. Ogden, and
H. Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Pomeroy, D.E., and M.H. Woodford. 1976. Drug
immobilization of Marabou Storks. J. Wildl.
M«na£.40:177-179.
Pooley, A.C. 1967a. Bird/Crocodile and bird/
hippopotamus commensalism in Zululand.
Ostrich 38:11-12.
Pooley, A.C. 1967b. Some miscellaneous
ornithological observations from the Ndumu
Game Reserve. Ostrich 38:31-32.
Pooley, A.C. 1968. Spoonbill and heron.
Lammergeyer 8:50.
Poonswad, P. 1979. Parasites of Asian Open-
billed Stork (Anastomus oscitans) and their
relationships to Pila and domestic duck. M.S.
Thesis, Mahidol University, Bangkok,
Thailand.
Poonswad, P., and P. Chatikavanij. 1989.
Echinoparyphium oscitansi N.sp. (Trematoda:
Echinostomatidae): natural infection in Asian
Open-billed Storks (Anastomus oscitans; Aves:
Ciconiidae) in Thailand. J. Sci. Soc. Thai.
15:293-299.
Poorter, E.P.R. 1969. Voedselzoekgewoonten en
voedselgebieden van de Lepelaars van het
Naardermeer. Rijksinst. veldbiol. Onderroek
Natuurb., Holland.
Poorter, E.P.R. 1979. Resultaten van het
onderzoek naar het voedsel en de
voedselgebieden van de lepelaars van het
Naardermeeer in 1970. Rijp Rapport, 1979-3
Abs. Rijksdienst voor de Ijsselmeerpolders, Lelystad.
66pp.
Poorter, E.P.R. 1982. Migration et dispersion des
Spatules neerlandaises. Oiseau Rev. Fr. Orn.
52:305-334.
Popa, G. 1989. Erfahrungen ueber
Schuelereinsatze zum Schutz des Weiss-
Storchs. Pp. 126-127 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Portela, L.I. 1986. A review of the status of the
Glossy Ibis Plegadis falcinellus in Portugal.
Cyanopica 4:693-712.
Porter, R. 1973. The disappearing ibis. Birds
(RSPB Mag.) 4:227-228.
Porter, R. 1974. The Bald Ibis or Waldrupp.
Bull. Orn. Soc. Turkey 8:4-5.
Porter, R., and I. Willis. 1968. The autumn
migration of soaring birds at the Bosphorus.
Ibis 110:520-536.
Portielje, A.J.F. 1927. Ornitho-psychologische
opmerkingen. Ardea 16:20-22.
Portnoy, J.W. 1978. A wading bird inventory of
coastal Louisiana (USA). Pp. 227-234 in
Sprunt, A., J.C. Ogden, and S. Winckler (eds),
Wading Birds. Research report 7, National
Audubon Society, New York.
Portnoy, J.W. 1981. Breeding abundance of
colonial waterbirds on the Louisiana-
Mississippi-Alabama coast. Am. Birds 35:868—
872.
Post, H., and F. Rueb. 1987. Bigamie bij
ooievaars (Ciconia ciconia). Het Vogeljaar
35:325-327.
Post, H., F. Rueb, and R. Belterman. 1991.
Sexing of White Storks (C. ciconia}. J. Orn.
132:99-101.
Post, P.W. 1962. Glossy Ibis breeding in New
York. Auk 79:120-121.
Post, W. 1990. Nest survival in a large ibis-heron
colony during a 3-year decline to extinction.
Colonial Waterbirds 13:50-61.
Post, W., F. Enders, and T.H. Davis, Jr. 1970.
The breeding status of the Glossy Ibis in New
York. Kingbird 20:3-8.
Poulsen, H. 1950. Beobachtungen an einem
Schuhschnabel (Balaeniceps rex) in der
Gefangenschaft. Z. Tierpsychol. 7:134-137.
Poulsen, H. 1951. Colour films: the greeting
ceremony of the Shoe-billed Heron, Balaeniceps
rex]. Gould, and the courtship display of the
Condor, Vultur gryphus L. Proc. Int. Orn. Congr.
10:396-397.
Powell, G.V.N. 1983. Food availability and
reproduction by Great White Herons, Ardea
herodias: a food addition study. Colonial
Waterbirds 6:139-147.
365
Powell, G.V.N. 1987. Habitat use by wading
birds in a subtropical estuary: implications of
hydrography. Auk 104:740-749.
Powell, G.V.N., R.D. Bjork, J.C. Ogden, R.T.
Paul, A.H. Powell, and W.B. Robertson, Jr.
1989. Population trends in some Florida Bay
wading birds. Wilson Bull. 101:436-457.
Powlesland, R., and D. Crockett. 1986. A Glossy
Ibis in the Chatham Islands. Notornis 33:232.
Prager, E.M., and A.C. Wilson. 1975. Slow
evolutionary loss of the potential for
interspecific hybridization in birds: a
manifestation of slow regulatory evolution.
Proc. Nat. Acad. Sci. USA. 72:200-204.
Prater, S.H. 1931. The migration of the White
Stork (Ciconia ciconia). J. Bombay Nat. Hist. Soc.
35:459.
Preston, F.W. 1969. Shapes of birds' eggs: extant
North American families. Auk 86:246-264.
Preston, I. 1976. Whale-headed Stork. Bull. E.
Afr. Nat. Hist. Soc. Nov/Dec:131-132.
Price, E.W. 1936. A new heterophyd trematode
of the genus Ascocotyle (Centrocestinae). Proc.
Helminthol. Soc. Wash. 3:31-32.
Priest, C.D. 1933. The Birds of Southern Rhodesia.
William Clowes, London.
Priest, C.D. 1948. Eggs of Birds Breeding in Southern
Africa. University Press, Glasgow.
Prigogine, A. 1953. Contribution 1'etude de la
fauna ornithologique de la region 1'ouest du
lac Edouard. Annal. du Musce Rogue Long Beige
Tervuren (Belg.), Ser. 8, Sci. Zool. 24.
Prigogine, A. 1976. Additions a 1'avifaune du
Zaire. Gerfaut 66:307-308.
Priklonskii, S.G. 1958. [Material on the ecology
of the Black Stork in the Oka Nature Reserve.]
Trud. Oksk. Gosudarstwennogo Sapowednika II.
Raboty Okskoi Orn. St. 1:102-115.
Priklonskii, S.G. 1977. [Numbers of Black Storks
in the Oka Forest Reserve.] P. 240 in Winter,
S. (ed.), VII All-Soviet Ornithological Conference.
Zoological Institute of the Academy of Science
of the Ukranian SSR, Kiev.
Priklonskii, S.G., and V.M. Galushin. 1959.
[New data on the ecology of the Black Stork.]
Proc. Baltic Orn. Conf. (1957) 3:231-236.
Prinzinger, R., and K. Hund. 1982. [Leg-
excreting in the White Stork (Ciconia ciconia) a
thermoregulatory behavior for body-cooling
through the legs.] Okel. Vogel 4:82-83.
Profus, B., Z. Jakubiec, and P. Miekczarek. 1989.
Zur situation des Weiss-Storchs in Polen.
Stand 1984. Pp. 81-98 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz. Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Prozesky, O.P.M. 1980. A Field Guide to the Birds
of Southern Africa. Collins, London.
Przhevalsky, N.M. 1870. A Journey Across
Ussurysky Territory, 1867-1869. St. Petersburg,
Russia.
Psenner, H. 1969. Der Innsbrucker Alpenzoo.
Gefoderte Welt 93:103.
Puerta, M.L., R. Munoz Pulido, V. Huecas, and
M. Abelenda. 1989. Hematology and blood
chemistry of chicks of White and Black Storks
(Ciconia ciconia and Ciconia nigra). Comp.
Biochem. Physiol. 94:201-204.
Pukinskij, Y.B. 1981. [Density and distribution
of rare and endangered species of birds in the
Primorye and Bikin River Valley.] Pp. 1-92 in
Bromlej, G.F., V.A. Kostenko, V.G. Yudin,
and V.A. Nechaev (eds.) Rare and Endangered
Terrestrial Animals of the Far East of the USSR.
Akad Nauka, USSR Vladistok.
Pundt, G., and H. Ringleben. 1963. Der Loeffler
(Platalea leucorodia) 1962 erstmals deutscher
Brutvogel auf der Insel Memmert. J. Orn.
104:97-100.
Purchase, D. 1976. The occurrence of Threskiornis
 molucca strictipennis in New Guinea. Emu
76(2):89.
Putnam, L., G. Maxwell, and S. Tilley. 1964.
Sight record of the Glossy Ibis for the Bass
Islands, Lake Erie, Ohio. Wilson Bull. 76:98.
Putzig, P. 1938. Zur Ernahrung des Weissen
Storch. Beitr. Fortpfanzungsbiol. Vogel 14:107-
108.
Pym, F.A.O. 1909. A list of the birds of the
Kaffrarian frontier. J. S. Afr. Orn. Union 5:91—
113.
Quay, T.L., and D.A. Adams. 1956. Nesting of
Cattle Egrets and Glossy Ibis in the Battery
Island Rookery at Southport, N.G. Chat 20:56-
57.
Quelch, JJ. t888. A collecting trip on the Abary
Creek. Timerhi 4:358-378.
Quelch, JJ. 1891. The native birds of
Georgetown. Timehri 5:69-107.
Queney, A. 1952. Cambriolage chez les cigognes.
Bull. Mens. Soc. Linn. Lyon 21:206-207.
Quickelberge, C.D. 1972. Spoonbills breeding in
the Eastern Cape Province. Ostrich 43:67.
Quinton, W.F. 1976. Letter to editor. Bokmakierie
28:76.
Rabor, D.S. 1977. Philippine Birds and Mammals.
Univ. Philippines Press, Quezon City.
Radkiewicz,J. 1989a. [Distribution and numbers
of White Stork Ciconia ciconia L. in Zielona
Gora district in 1985.] WSP-Zielona Gora
Przyr. Strodk. Nad. 1:47-66.
Radkiewicz, J. 1989b. Der WeiBstorch im
mittleren Westpolen 1984. Pp. 99-104 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz, Proc. 1st Int.
Stork Conserv. Symp. Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Radkiewicz, J., and E. Schuz. Gibt es Bigamie
beim WeiBstorch? Orn. Mitt. 38:139.
Radolfzell, V. 1950. Der Storchbestand in
Sudwestdeutschland 1949. Rundbrief 1950 der
Vogelwarte Radolfzell.
Raffaele, H.A. 1983. A Guide to the Birds of Puerto
Rico and the Virgin Islands. Fondo Educ.
Interamer, San Juan, Puerto Rico.
Rahmani, A.R. 1987. Is the Blacknecked Stork
threatened? Hornbill 1987(4): 18-19.
Rahmani, A.R. 1988. Blacknecked Stork.
Newsletter Birdwatchers 27(3/4):8-9.
Rahmani, A.R. 1989a. Status of the Blacknecked
Stork Ephippiorhynchus asiaticus in the Indian
subcontinent. Forktail 5:99-110.
Rahmani, A.R. 1989b. Greater Adjutant Stork.
Newsletter Birdwatchers 29(ll&12):2-3.
Rahmani, A.R. 1989c. Blacknecked and Greater
Adjutant storks in India. ICBPIIWRB Spec.
Group Storks, Ibis, Spoonbills Newsletter 2:3-6.
Rahmani, A. 1991. Form for surveying storks in
Indian zoological gardens. Zoos' Print 6(l):5-6.
Rahmani, A.R., G. Narayan, and L. Rosalind.
1990. Status of the Greater Adjutant Leptoptilos
dubius in the Indian subcontinent. Colonial
Waterbirds 13:139-142.
Raines, RJ. 1962. The distribution of birds in
northeast Greece in summer. Ibis 104:490-502.
Rallo, G. 1975. Capture of rare and infrequent
birds in the Venezia Province Italy.
Ornithology of Venezia Part 3. Boll. Mus. Civ.
Stor.Nat. Ven. 27:15-20.
Ramo, C., and B. Busto. 1982. Son Eudocimus
ruber y E. albus distintas especies? Dohana Ada
Vertebr. 9:404-408.
Ramo, C., and B. Busto. 1984. Censo aereo de
Corocoros (Eudocimus ruber} y otras aves
acuaticos en Venezuela. Bol. Soc. Venezolana
Cienc. Nat. 39:65-88.
Ramo, C., and B. Busto. 1985. Compartamiento
reproductive del corocoro (Eudocimus ruber) en
los llanos de Venezuela. Mem. Soc. Cien. Nat. la
Salle 45:77-113.
Ramo, C., and B. Busto. 1987. Hybridization
between the Scarlet Ibis (Eudocimus ruber) and
the White Ibis (E. albus) in Venezuela. Colonial
Waterbirds 10:111-114.
Ramo, C., and B. Busto. 1988. Status of the
nesting population of the Scarlet Ibis
(Eudocimus ruber) in the Venezuelan llanos.
Colonial Waterbirds 53:311-314.
Ramo, C., and B. Busto. no date. El Corocoro Rojo
y su Mundo. Med. Edit., C.A., Caracas.
Ramo, C., and B. Busto. in press. Los garceros
del area Guanare-Masparro. Mem. Soc. Hist.
Nat. la Salle.
Ramo, C., E. Aguilera, B. Busto, and S. Reid.
1983. Censo aereo preliminar de las colonias
de cria de Eudocimus ruber y otras aves acuaticas
de los Llanos de Venezuela. Rev. Univ. Nac.
Exp. Llanos Occident. Ezequiel Zamorra Cienc.
Tecnol. 1:61-69.
Ramsamujh, B. 1990. Status and conservation of
the Scarlet Ibis in Guyana. Pp. 95-99 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11.
IWRB, Slimbridge, UK.
Ramsden, C.T. 1913. Glossy Ibis (Plegadis
autumnalis) in eastern Cuba; a new record. Auk
30:268.
Ramsey, JJ. 1968. Roseate Spoonbill chick
attacked by ants. Auk 85:325.
Rana, B.D. 1988. Migratory birds in the desert
region of Rajasthan. Tigerpaper 15:29-31.
Rand, A.L. 1936. The distribution and habits of
Madagascar birds. Bull. Am. Mus. Nat. Hist.
72:143-499.
Rand, A.L. 1956. Foot-stirring as a feeding habit
of Wood Ibis and other birds. Am. Midi. Nat.
55:96-100.
Rand, A.L., and R.L. Fleming. 1957. Birds from
Nepal. Fieldiana Zoo/. 41:1-218.
Rand, A.L., and E.T. Gilliard. 1967. Handbook of
New Guinea Birds. Weidenfeld and Nicolson,
London.
Randik, A.K. 1989. A summary of habitat
changes and their effect on breeding
populations of the White Stork in the
Carpathian Basin, C.S.S.R. Pp. 403-404 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Ranner, A. 1989. Entwicklung und derzeitige
Situation des Weiss-Storchs in Rust. Vogelsch.
Osterreich 4:29-33.
Rao, P., and S. Murlidharan. 1989. Unusual
feeding behaviour of the Adjutant Stork,
Leptoptilos dubius (Gmelin). J. Bombay Nat. Hist.
Soc. 86:97.
Rao, P. and S. Murlidharan. 1990. Weight of
Whitenecked Stork Ciconia episcopus.J. Bombay
Nat. Hist. Soc. 87:139.
Rasajski, J. 1989. [Numbers of nesting pairs of
White Storks C. ciconia L. and associated
occurrences of nesting in south Banat region,
1976-1985.] Z,<mw 40:111-123.
Raseroka, B.H. 1975a. Breeding of the Hadedah
Ibis. Ostrich 46:208-212.
Raseroka, B.H. 1975b. Diet of the Hadedah Ibis.
Ostrich 46:51-54.
Rasmussen, D.T., S.L. Olson, and E.L. Simons.
1987. Fossil birds from the Oligocene Jebel
Qatrani Formation, Fayum Province, Egypt.
Smithson. Contrib. Paleobiol. 62. 20pp.
Rasmussen, E.V. 1979. Den sorte storks (Ciconia
nigra) forekomst i Danmark, isaer i arene
1970-1978. [The occurrence of the Black Stork
(Ciconia nigra) in Denmark, 1970-1978.] Dan.
Orn. Foren. Tidsskr. 73(4):265-270.
Rattey, F. 1965. Am Horst des Schwarzstorchs.
Falke 12:245.
Rauber, E.L., and D. Dowdy. 1972. Roseate
366
Spoonbill identified at Tennessee National
Wildlife Refuge. Migrant 43:67.
Rauhe, H. 1955. Gehauftes Auftreten des
Weissen Storches (C.c.) zwischen Elbe—und
Wesermundung. Ornithol. Mitt. 7:91.
Raviv, M. 1989. Ring recoveries of the White
Stork in Israel and Sinai, 1909-1982. Pp. 317-
324 in Rheinwald, G., J. Ogden, and H.
Schulz (eds.), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Reader's Digest Association. 1974. The Book of
British Birds, 2nd edn. Reader's Digest
Association. London.
Rechnitzer, A.B. 1954. Status of the Wood Ibis
in San Diego County, California. Condor
56:309-310.
Rechnitzer, A.B. 1956. Foraging habits and local
movements of the Wood Ibis in San Diego
County, California. Condor 58:427-432.
Ree, V. 1973. Dagens avifaunistiske situasjon i
Las Marismas i Sor-Spania. Sterna 12:225-268.
Reed, C.A., and T.E. Lovejoy. 1969. The
migration of the White Stork in Egypt and
adjacent areas. Condor 71:146-154.
Reed, C.K. 1901. White Ibis. Am. Orn. 1(3):38-
39.
Reed, S. 1980. Glossy Ibis among South Island
pied oystercatchers. Notornis 27(2): 124.
Reichenbach, H.G.L. 1849-1852. AviumSystema
Naturale, Aves Grallatores. Hofmeister, Dresden
and Leipzig.
Reichenow, A. 1877. Systematische Uebersicht
der Schreitvogel (Gressores), einer
naturlichen, die Ibidae, Ciconiidae,
Phoenicopteridae, Scopidae, Balaenicipidae
und Ardeidae umfassenden Ordnung.J. Orn.
1877:113-171,229-277.
Reichenow, A. 1900-1901. Die Voegel Afrikas, vol.
1. Neudmann, Berlin.
Reid, G. 1881. The birds of the Lucknow Civil
Division, part 2. Stray Feathers 10:1-88.
Reilly, T.E., and B. Wasdell. 1965. Marabou
stork Leptoptilos crumeniferus (Lesson) breeding
in Swaziland. Ostrich 36:96.
Reimann, EJ. 1939. Banding adult Roseate
Spoonbills. Oologist 56:88-90.
Reinhardt, J. 1860. On the affinities of
Balaeniceps. Proc. Zoo/. Soc. Lond. 1860:377-380.
Reinhardt, J. 1862. Some remarks on the genus
Balaeniceps. /£w 4:158-175.
Reither, H. 1986. Der Weiss-Storch (Ciconia
ciconias) im Verwaltungsbezirk Braunschweig
1985 und 1986- ein Vergleich. Beitr. Naturk.
Niedersachsens 39:245-246.
Reither, H. 1987. Ubersicht uber den Brutertfolg
des WeiBstorches (Ciconia ciconia) im
Regierungsbezirk Braunschweig 1987. Beitr.
Naturk. Niedersachsens 41:114.
Rejman, B. 1989. Der WeiBstorchs in der
Tschechei 1934-1984. Pp. 69-72 in Rheinwald,
G., J. Ogden, and H. Schulz (eds), Weiss-
Storch: Status undSchutz, Proc. 1st Int. Stork
Conserv. Symp., Walsrode, 1985, Schriftenr.
DDA, Braunschweig, Germany.
Rejman, B. and C. Folk. 1991. Results of the 4th
International White Stork census in the Czech
Republic. Fol. Zoo/. 40:179-186.
Rekasi, J. 1989. Nahrungsbiologische
Untersuchungen am WeiBstorch. Pp. 397-402
in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weiss-Storch: Status und Schutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Remsen, J.V. 1986. Aves de una localidad en la
sabana humeda del norte de Bolivia. Ecol.
Bolivia 8:21-35.
Remsen, J. v., M.A. Traylor, Jr., and K.C.
Parkes. 1985. Range extensions for some
Bolivian birds, 1 (Tinamiformes to
 Charadriiformes). Bull Br. Orn. Club 105:124-
130.
Remy, J.M. 1956. Observations sur les vols de
cigognes au Sahara Central. Oiseau Rev. Fr.
Orn. 26:152-153.
Rencurel, P. 1974. L'Ibis chauve Geronticus
eremita dans le Moyen-Atlas. Alauda 42:143-
158.
Rengstorf, D t 1990. A White-faced Ibis in
Dakota and Hennepin Counties. Loon 62:154-
156.
Renjifo, S. 1950. Contribuciones a la
parasitologia Columbiana. II. Haemoparasitos
de aves y otrosvertebrados de low Llanos
Orientales. Rev. Acad. Colomb. Cienc. 7:539-547.
Renjifo, S., C. San Martin, and J de Zulueta.
1952. A survey of the blood parasites of
vertebrates in eastern Columbia. Acta Trop.
9:151-169.
Rensch, B. 1931. Die Vogelwelt von Lombok,
Sumbawa, und Flores. Mitt. Zoo/. Mus. BerL
17:451-637.
Reygaert. 1982. De doortrek van de Zwarte
Ooievaar (Ciconia nigra) in Belgie t.m. 1981.
Ornis Flandriae 1982:89-99.
Reynolds, J. 1965. Association between Little
Egret and African Spoonbill. Birds 58:468.
Reynolds, J. 1976. Africa's striking storks.
Wildlife (Lond.) 18:347-351.
Reynolds, J.F. 1970. Storks wing spreading while
feeding. British Birds 70:4.
Reynolds, J.F. 1972. Some new records for
Tanzania National Parks. Bull E. Afr. Nat.
Hist.Soc. 1972:141.
Reynolds, J.F. 1977. Thermo-regulatory
problems of birds nesting in arid areas in East
Africa: a review. Scopus 1:57-68.
Rheinwald, G. 1989a.
Wiedereinbuergerungsprogramme fur den
Weiss-Storch. Einleitung. [Reintroduction.
Introduction.] Pp. 435-436 in Rheinwald, G.,
J. Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Rheinwald, G. 1989b. Versuch einer Bilanz. Pp.
221-227 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Rheinwald, G. 1989c. Brut: Bestand, Trends,
und Biologic. Einfuehrung. Pp. 17-18 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status und Schutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Rheinwald, G. 1989d. Biologic und Schutz:
Einfuhrung. Pp. 379-380 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Rheinwald, G. 1989e. Nachwort [Epilogue]. Pp.
468-471 in Rheinwald, G., J. Ogden, and H.
Schulz (eds), Weiss-Storch: Status und Schutz,
Proc. 1st Int. Stork Conserv. Symp., Walsrode,
1985, Schriftenr. DDA, Braunschweig,
Germany.
Rheinwald, G., J. Ogden, and H. Schulz (eds)
1989. Weiss-Storch: Status und Schutz [White Stork :
Status and Conservation], Proc 1st Int. Stork
Conserv. Symp., Walsrode 1985, Schriftener.
DDA, Braunschweig, Germany. 472pp.
Rice, D.W. 1955. The Glossy Ibis of Lake Alice.
Q.J. Fla. Acad. Sci. 18(1):20.
Rich, P.V. 1972. A fossil avifauna from the
Upper Miocene Beglia Formation of Tunisia.
NotesduServ. Geol. (Tunis.) 35:29-66.
Richard, E., and C. Laredo. 1988. AJabiru
mycteria record outside its usual range,.
Mendoza Prov. (Argentina). Nuestras Aves Bol.
Asoc. Orn. del Plata 15:12-13.
Rickett, C.B. 1903. Additional field-notes on the
birds of Fohkien. Ibis (ser. 8) 10:215-222.
Ricklefs, R.E. 1968. Patterns of growth in birds.
Ibis 110:419-451.
Ridgely, R.S. 1976. A Guide to the Birds of Panama.
Princeton University Press. Princeton, New
Jersey.
Ridgway, R. 1874. Notes upon American water
birds. Am. Nat. 8:108-111.
Ridgway, R. 1878. Studies of the American
Herodiones. Part 1—Synopsis of the American
genera of Ardeidae and Ciconiidae; including
descriptions of three new genera, and a
monograph of the American species of the
genus Ardea, Linn. US Geol. Geog. Surv. Terr.
4(1):219-251.
Ridgway, R. 1884. Remarks upon the close
relationship between the White and Scarlet
Ibises Eudocimus albus & Eudocimus ruber. Auk
1:239-240.
Ridgway, R. 1893. Description of some new birds
collected on the islands of Aldabra and
Assumption northwest of Madagascar by Dr.
W. L. Abbott. Proc. US Natl Mus. 16:599.
Ridgway, R. 1895. On birds collected by Doctor
W.L. Abbott in the Seychelles, Amirantes,
Gloriosa, Assumption, Aldabra, and adjacent
islands, with notes on habits, etc., by the
collector. Proc. U.S. Nat. Mus. 18(1079):509-
546.
Riegel, M., and W. Winkel. 1971. Ueber
Todesursachen beim WeiBstorch (C. ciconia}
and Hand von Ringfundangaben. Vogelwarte
26:128-135.
Rijk, J., de. 1989. [The distribution of the White
Stork C. ciconia in the Netherlands in the 18th
century.] Limosa 62:63-66.
Riley, J.H. 1925. A collection of birds from north
and north-central Celebes. Proc. US Natl Mus.
64:1-118.
Riley, J.H. 1938. Birds from Siam and the Malay
Peninsula in the United States National
Museum. US Natl Mus. Bull. 172:1-581.
Riley, L. 1981. The stately, handsome stork. Bird
Watcher's Dig. Jul/Aug:40-46.
Ringleben, H. 1936. Uber einige Falle von
Polyandrie und Polygamie bei Voegeln. Orn.
Monatsber. 44:178-179.
Ringleben, H. 1940. Uber den Eintritt der
Geschlechtsreife und liber den Aufenthalt
nichtbrutender Voegel, insbesondere Strand-
und Seevoegel, zur Brutzeit. Beitr.
Fortpflanzungsbiologie d. Voegel 16:10-23.
Ringleben, H. 1950. Zur Ausbreitung und
Verbreitung der WeiBstorch in Nord-
osteuropa. Orn. Ber. 3:27-53.
Ringleben, H. 1986. An unnoticed stork colony
Ciconia ciconia in a village in Brandenburg, East
Germany. Beitr. Vogelkd. 32:331.
Rios U.G. 1990. Conservacion de la fauna
silvestre por iniciativa privada en los llanos
occidentales. Projecto de cooperacion mutua.
Pp. 85-94 in Frederick, P.C., L.G. Morales,
A.L. Spaans, and C.S. Luthin (eds), The Scarlet
Ibis (Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11.
IWRB, Slimbridge, UK.
Rios, G., and G. Morales (compilers). 1990.
Estado de las poblaciones del Corocoro Rojo
(Eudocimus ruber) en Venezuela: diagnostico,
conclusiones y recomendaciones para un plan
de conservacion. Pp. 137-151 in Frederick,
P.C., L.G. Morales, A.L. Spaans, and C.S.
Luthin (eds), The Scarlet Ibis (Eudocimus
ruber): Status, Conservation, and Recent Research,
Proc. 1st International Scarlet Ibis
Conservation Workshop, Caracas, Venezuela.
367
IWRB Special Publication 11, Slimbridge,
Glos, UK.
Ripley, S.D. 1961. A Synopsis of the Birds of India
and Pakistan Together with those of Nepal, Sikkim,
Bhutan and Ceylon. The Bombay Natural
History Society, Bombay. 703pp.
Ripley, S.D. 1963. Brief bird observations in
Nubia. Ibis 105:108-109.
Risdon, D.H.S. 1969. The breeding of Scarlet
Ibis Eudocimus ruber. Avic. Mag. 75:165-167.
Risdon, D.H.S. 1971. Breeding the Sacred Ibis
and the Scarlet Ibis Threskiornis aethiopica and
Eudocimus ruber at the Tropical Bird Gardens,
Rode. Int. Zoo Yb. 11:131-132.
Ritzel, L. 1980. Aquila—Adler attackiert
Schwarzstorch (Ciconia nigra). Orn. Mitt.
32:23-24.
Robbins, C.S., B. Bruun, and H.S. Zim. 1966. A
Guide to Field Identification of the Birds of North
America. Golden Press, New York.
Robert, H.C., J.M. Teal, and E.P. Odum. 1956.
Summer birds of Sapelo Island Georgia: a
preliminary list. Oriole 21:37-45.
Roberts, A. 1905. A visit to a breeding colony of
Ibis aethiopica (Sacred Ibis).J. S. Afr. Orn.
Union 5:32-33.
Roberts, A. 1932. Migration of African birds.
Ostrich 3:97-109.
Roberts, A. 1935. Scientific results of the Vernay-
Lang Kalahari Expedition, March to
September, 1930. Ann. Transvaal Mus. 16:
1-185.
Roberts, A. 1940. The Birds of South Africa. H.F.
and G. Witherby, London.
Roberts, A. 194 la. Notes on some birds of the
Cape Province. Ostrich 11:112-135.
Roberts, A. 1941b. The White Stork in South
Africa. Ostrich 12:31-41.
Roberts, A. 1942. More about the breeding of
White Storks in South Africa. Ostrich 13:17-24.
Roberts, TJ. 1969. A note on Ciconia nigra
(Linnaeus) in West Pakistan. J. Bombay Nat.
Hist. Soc. 66:616-619.
Robertson, D. 1980. Rare Birds of the West Coast of
North America. Woodcock Publishing, Pacific
Grove, California.
Robertson, H.A., and B.E. Preece. 1980. Juvenile
Royal Spoonbills Platalea regia at the
Manawatu Estuary, New Zealand. Notornis
27:170-171.
Robertson, W.B. 1953. More eastern Glossy
Ibises in Everglades National Park. Everglades
Nat. Hist. 1(3): 127-128.
Robertson, W.B., Jr. 1962. Ornithology of The
Cruise of the Bonton'. Tequesta XXII:65-77.
Robertson, W.B., Jr., and J.A. Kushlan. 1974.
The southern Florida avifauna. Pp. 414-452 in
Gleason. PJ. (ed.), Environments of South
Florida: Present and Past. Miami Geological
Society, Miami.
Robertson, W.B., Jr., and H.B. Muller. 1961.
Wild winds and wildlife. Audubon Mag. Nov-
Dec:308-311.
Robertson, W.B., and D.R. Paulson. 1961.
Florida region. Audubon Field Notes 15(1):
3-12.
Robertson, W.B., Jr., L.L. Breen, and B.W.
Patty. 1983. Movement of marked Roseate
Spoonbills in Florida with a review of present
distribution./ Field Orn. 54:225-236.
Robiller, F., and K. Trogisch. 1982. Ueber die
Schwarzstoerche, Ciconia nigra, im Vogelpark
Walsrode. Die Europ. Vogel. 2:28-29.
Robiller, F., and K. Trogisch. 1986a. Ueber den
Schwarzstorch und seine Zucht im Vogelpark.
Gefederte Welt 110(7): 189-191.
Robiller, F., and K. Trogisch. 1986b. Haltung
und Zucht von Loefflern im Vogelpark.
Gefaderte Welt 110:266-269.
Robin, P. 1958. Ibis a tete chauve, Comatibis
eremita, centre de nidification de Tasdrem.
C. R. Soc. Sci. Nat. Phys. Maroc 24:33-34.
Robin, P. 1973. Comportement des colonies des
Geronticus eremita dans le sud marocain, lors des
periodes de secheresse. Bonn Zoo/. Beitr.
24:317-322.
Robinson, G.G.W., and R.A. Ryder. 1971.
Snowy Egret, Black-crowned Night Heron and
White-faced Ibis banding in Colorado. Colo,
Field Orn. 9:1.
Robinson, H.C., and C.B. Kloss. 1911. On birds
from the northern portion of the Malay
Peninsular, including the islands of Lang Kawi
and Terstau, with notes on other rarer
Malayan species from the southern districts.
7^(9)5:10-80.
Robinson, H.C., and F.N. Chasen. 1936. The
Birds of the Malay Peninsula, vol. 3. H. F. and G.
Witherby, London.
Robson, C. 1985a. Recent reports—China.
Oriental Bird Club Bull. 2:38.
Robson, C. 1985b. Recent reports: Indonesia.
Oriental Bird Club Bull. 1:27-28.
Rockingham-Gill, D.V., and PJ. Mundy. 1989.
Status and distribution of the White Stork
(Ciconia ciconia) in Zimbabwe. Pp. 361-364 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriflenr. DOA, Braunschweig, Germany.
Rodgers, J.A. Jr. 1976. Spread-wing sunbathing
by juvenile White-faced Ibis. Auk 93:375-376.
Rodgers, J.A. Jr. 1978. Feeding association
between Brown Pelicans and Wood Storks.
Fla. Field Nat. 6:44-45.
Rodgers, J.A. Jr. 1980. Breeding ecology of the
Little Blue Heron on the west coast of Florida.
Condor 82:164-169.
Rodgers, J.A. Jr. 1986. A field technique for
color-dyeing nestling wading birds without
capture. Wildl. Soc. Bull. 14:399-400.
Rodgers, J.A. Jr. 1989. Minimum buffer zone
requirements to protect nesting colonial
waterbirds from human disturbance. Colonial
WaterbirdSoc. Newsletter 13(2): 17 (abstr.)
Rodgers, J.A. Jr. 1990. Breeding chronology and
clutch information for the Wood Stork from
museum collections. J. Field Orn. 61:47-53.
Rodgers, J.A., Jr., and D. Maeher. 1982. Wading
birds and wetlands. Florida Wildl. Jul—Aug
1982:30-33.
Rodgers, J.A., Jr., and S.A. Nesbitt. 1980.
Feeding energetics of herons and ibises at
breeding colonies. Proc. 1979 Colonial Waterbird
Conf. 3:128-132.
Rodgers, J.A., Jr., A.S. Wenner, and S.T.
Schwikert. 1987. Population dynamics of
Wood Storks in north and central Florida.
Colonial Waterbirds 10:151-156.
Rodgers, J.A., Jr., A.S. Wenner, and S.T.
Schwikert. 1988. The use and function of green
nest material by Wood Storks. Wilson Bull.
100:411-423.
Rogers, T.H. 1974. The spring migration April
1-May 31, 1974. Northern Rocky Mountain-
Intermountain region. Am. Birds. 28:828-832.
Rohl, E. 1942. Fauna Descriptiva de Venezuala.
Caracas.
Rohr, W. 1969. Waldrapp im Tiergarten
Heidelberg gezuchtet. Gefaderte Welt 93:182-
183.
Rome, A., and G. Travison. 1982. The marsh of
Castiglione Delia Pescaia, Italy. Report for
1981. Riv. Ital. Orn. 52:218-220.
Romero, A. 1983. Distress call saves a Caiman c.
crocodilus hatchling in the Venezuelan Llanos.
Biotropica 15(1):71.
Roosevelt, T. 1910. African Game Trails.
Scribners, New York.
Roosevelt, T. 1914. Through the Brazilian
Wilderness. Scribners, New York. 383pp.
Root, A. 1963. Notes on the feeding habits of the
Openbill Stork Anastomus lamelligerus. Ibis
105:399-400.
Rooth, J. 1957. Over het voedsel, de terreinkeus
en de achteruitgang van de Ooievaar, Ciconia
ciconia L., in Nederland Ardea 45:93-116.
Rooth, J. 1966. Vogeltelling in het hele
Nederlandse Waddengebied Augustus 1963.
Limosa 39:175-181.
Rooth, J., and D.A. Jonkers. 1972. The status of
some piscivorous birds in the Netherlands.
T. N. O.-Nieuws 27:551-555.
Roozendaal, D. 1988. Waarnemingen aan de
paarvorming van de Rode Ibis (Eudocimus
ruber) in de dierentuin Artis, Amsterdam.
Afdeling Diergedrag, Universiteit van
Amsterdam, doctoraal verslag.
Roscoe, EJ. 1955. Aquatic snails found attached
to feathers of White-faced Glossy ibis. Wilson
Bull. 67(1):66.
Rose, A.B. 1973. Food of some Australian birds.
Emu 73:177-183.
Rosenberg, E. 1941. Vid svarta storkens bo.
Fauna Flora 36:1-10.
Rosenberg, H., von 1887. Wie alt ein Storch
werdenkann. Mitt. Orn. Ver. Wien, 11:64.
Roseveare, M. 1989. First record of the Shoebill
in Malawi. Nyala 14:45.
Roslyakov, G.E. 1977. [Rare birds in the lower
Priamura Region.] Pp. 241-243 in Winter, S.
(ed.), VII All-Soviet Ornithological Conference.
Zoological Institute of the Academy of Science
of the Ukranian SSR, Kiev.
Rossbach, R. 1978. Zum Bestand des Weiss-
Storchs (Ciconia ciconia) in Hessen (1975-
1978). [On the status of white storks (Ciconia
ciconia) in Hesen (1975-1978).] Lucinia43(5/
6):182-186.
Roth, W.E. (ed.). 1922-1923. RichardSchomburgk's
Travels in British Guyana 1840-44, vol. 1.
Georgetown, Guiana.
Rothschild, W., E. Hartert, and O.
Kleunschmidt. 1897. Comatibis eremita (Linn.)
A European bird. Novit. Zoo/. 4:371-377.
Round, P.D., B. Amget, J. Jintanugol, and U.
Treesucon. 1988. A summary of the larger
waterbirds in Thailand. Tiger Paper 15(3): 1-9.
Roux, F. 1962. Sight-records of ringed birds and
the use of national coloured supplementary
rings. Ring Int. Orn. Bull. 30-31:84-86.
Roux, F. 1972. West Africa Survey. Bull. Int.
Waterfowl Res. Bur. 34:33-36.
Roux, F. 1973. Recensement d'oiseaux
aquatiques dans le delta du Senegal. Oiseau
Rev.Fr. Orn. 43:1-15.
Rozdina, O.I., E.I. Sotnikova, and E.N.
Skakunova. 1991. Notes on Oriental White
Storks, Ciconia bqyciana, in the Moscow Zoo,
1985-88. Pp. 203-206 in Coulter, M.C., Q.
Wang, and C.S. Luthin. (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Rubio Garcia, J.C., M.R. de los Santos, and R.
Santa Rosa. 1983. [The breeding of White
Storks (Ciconia ciconia) of the 'marismas', the
Guadalquivir River (Spain).] Alauda 51:251-
258.
Ruckdeschel, C., and C.R. Shoop. 1987 (1989).
Aspects of Wood Stork nesting ecology on
Cumberland Island, Georgia. Oriole 52:1-27,
71.
Rudegeair, T. 1975a. The gular pouch of the
female White Ibis. Auk 92:168-169.
Rudegeair, T. 1975b. The reproductive behavior
and ecology of the White Ibis Eudocimus albus.
PhD Thesis, University of Florida, Gainsville,
Florida.
Ruppell, E. 1835-1840. Neue Wirbelthiere zu der
Fauna von Abyssinien gehb'rig. Frankfurt am
Main.
Ruppell, E. 1845. Systematische Uebersicht der Vogel
Nord-Ost-Afrika's. Frankfurt am Main.
Ruppell, G. 1977. Quantitative Unterschiede bei
368
Landemanovern von alten und jungen Ibissen
(Threskiornisaethiopicus).f. Orn. 118:282-287.
Ruppell, W. 1939. Klimaperioden als Ursache
fur die Zunahme der Stoerche. Deutsches
Weidwerk 1:7-9.
Ruppell, W. 1942. Versuch einer Storchzugkarte.
Vogelzug 13:35-39.
Russell, F. 1972. The coming of the ibis. Audubon
Mag. 74(6):52-55.
Russell, J.K. 1978. Effects of interspecific
dominance among egrets commensally
following Roseate Spoonbills. Auk 95:608-610.
Russell, R.P., Jr. 1982. Roseate Spoonbills feed
on vegetable material. Fla. Field Nat. 10(1):18.
Russell, S.M. 1964. A distributional study of birds of
British Honduras. Ornithological Monographs 1.
American Ornithologists' Union, Washington
D.C.
Rutgers, A. 1969. Birds of Asia. Taplinger
Publishing Company. New York, New York.
Rutgers, A., and K.A. Norris. 1970. Encyclopedia
of Aviculture, vol. 1. Blandford Press, London.
350pp.
Ruthke, P. 1957. Aus der pommerschen
Vogelwelt. II. Vom Schwarzstorch. Vogelwelt
78:157-162.
Ruthke, P. 1966. Beitrag zur Vogelfauna
Marokkos. Bonn. Zoo/. Beitr. 17:186-201.
Ruthke, P. 1977. Beobachtungen an
Jugendtrupps des Waldrapps (Eudocimus
eremita). Vogelwelt 98:231-233.
Ruwet, J.-C. 1963. Notes ecologiques et
ethologiques sur les oiseaux des plaines de la
Lufira superiure (Katanga). 1. Podicipedes,
Steganapodes, Gressores, Anseres. Rev. Zoo/.
Bot.Afr. 68:1-60.
Ryan, P.G., J. Cooper, CJ. Stutterheim, and R.
Loutit. 1984. An annotated list of the birds of
the Skeleton Coast Park. Madoqua 14:79-90.
Ryder, J.H., and B.A. Ryder. 1978. First
breeding records of Black Stork in Malawi.
Ostrich 49:51.
Ryder, R.A. 1967. Distribution, migration and
mortality of the White-faced Ibis (Plegadis
chihi) in North America. Bird-Banding 38:257-
277.
Rydzewski, W. 1962. Longevity of ringed birds.
Ring 3:147-152.
Rydzewski, W. 1973. Longevity records, III.
Ring 76:63-70.
Rydzewski, W. 1976. Longevity records X. Ring
86-87:17-18.
Rysavy, B., and J.K. Macko. 1973. Bird cestodes of
Cuba, vol. 1. Anales del Institute de Biologia.
Universidad Nacional Autonoma de Mexico
42 (Serie Zoologia 1971): 1-28.
Ryser, F.A., Jr. 1985. Birds of the Great Basin, a
Natural History. University of Nevada Press.
Reno, Nevada.
Sackl, P. 1985. Der Schwarzstorch (Ciconia nigra)
in Osterreich—Arealausweitung,
Bestandsentwicklung und Verbreitung.
Vogelwelt 106:121-141.
Sackl, P. 1985. Die Ausbreitung des
Schwarzstorchs (Ciconia nigra, Aves) —
Arealveranderungen in Osteuropa und Sudost-
Oesterreich. Mitt. Naturwiss. Ver. Steiermark
115:125-131.
Sackl, P. 1987. Ueber saisonale und regionale
Unterschiede in der Ernahrung und
Nahrungswahl des Weiss-Storches (Ciconia c.
ciconia) im Verlauf der Brutperiode. Egretta
30:49-80.
Sackl, P. 1989. Zur Ernahrungsbiologie und
Habitatnutzung des Weiss-Storchs. Vogelschutz
Osterreich 4:6-10.
Saes, H. 1967. Rhodotomla vuilleminii new species
isolated on a Red Ibis Guara rubra. Bull.
Trimest. Soc. Mycol Fr. 83:453-458.
Safriel, U. 1968. Bird migration at Elat, Israel.
7^110:283-320.
Safriel, U.N. 1980. Notes on the extinct
 population of the Bald Ibis Geronticus eremita in
the Syrian Desert. Ibis 122:82-88.
Sahin, R. 1980. Erfolgreiche Volierenbrut der
Waldrappe.in der Turkei. Orn. Mitt. 32:72-74.
Sahin, R. 1981. Breeding the Hermit Ibis
Geronticus eremita in Birecik, Turkey. Commun.
Fac. Sci. Univ. Ankara. Ser. C H, Zoo/. 25:37-44.
Sahin, R. 1982a. Eltern-Kind Beziehungen der
freilebenden Waldrappe (Geronticus eremita L.)
in Birecik (Turkei). Oekol. Voegel4:\-7.
Sahin, R. 1982b. Beitragzum
Fortpflanzungsverhalten der freilebenden
Waldrappe (Geronticus eremita L.) in der Turkei.
1. Mitteilung: Ankunft, Paarbildung und
Nisten. Oekol. Voegel 4:181-190.
Sahin, R. 1982c. Zur Form der Ehe Freilebender
Waldrappe (Geronticus eremita) in Birecik
(Turkei). Orn. Mitt. 34:162-163.
Sahin, R. 1983a. Beitrag zum
Fortpflanzungsverhalten der freilebenden
Waldrappe (Geronticus eremita L.) in der Turkei.
2. Mitteilung: Paarung. Oekol. Voegel 5:63-72.
Sahin, R. 1983b. Beitragzum
Fortpflanzungsverhalten der freilebenden
Waldrappe (Geronticus eremita L.) in der Turkei.
3. Mitteilung: Eiablage, Bruten, und
Schlupfen. Oekol. Voegel 5:152-155.
Sahin, R. 1983c. Beitragzum
Fortpflanzungsverhalten der freilebenden
Waldrappe (Geronticus eremita L.) in der Turkei.
4. Mitteilung: Fortpflanzungskampfe. Oekol.
Voegel 5:263-270.
Sahin, R. 1983d. Koerper-und Nesthygiene der
freilebenden Waldrappe (Geronticus eremita).
Orn. Mitt. 35:152-155.
Sahin, R. 1990. [Head-nodding in Bald Ibises
(Geronticuseremita).} J. Orn. 131:445-451.
Saiff, E.I. 1978. The middle ear of the skull of
birds: the Pelecaniformes and Ciconiiformes.
Zool. J.Linn.Soc. 63:315-370.
Saikia, P., and P.C. Bhattacharjee. 1989a. A
preliminary survey of Adjutant Storks in
Assam. Asian Wetland News 2(2):\4-\5.
Saikia, P., and P.C. Bhattacharjee. 1989b.
Adjutant storks at risk in Assam, India. Spec.
Group Storks, Ibis, Spoonbills Newsletter 2:6-8.
Saikia, P., and P.C. Bhattacharjee. 1990a.
Conservation of large wetland birds in Assam:
Adjutant Storks. Pp. 50-51 abstr. in Seminar on
Wetland Ecology and Management. Bombay Nat.
Hist. Soc., Bharatpur.
Saikia, P., and P.C. Bhattacharjee. 1990b.
Nesting records of Greater Adjutant Storks in
Assam, India. Spec. Group Storks, Ibis, Spoonbills
Newsletter 3(l/2):2-3.
Saikia, P., and P.C. Bhattacharjee. 1991. Status
of Ibises and Spoonbills in Assam. Spec. Group
Storks, Ibises, Spoonbills Newsletter 4(1):3.
Sakamoto, M. 1966. Kohnotori: Japanese Stork,
Ciconia ciconia boyciana (Swinhoe). Kobe
Newspaper Press, Kobe. 104pp.
Sakou, T., et al. 1978. The breeding of the
spoonbill. Animals and Zoos 30:294-297.
Salathe, T. 1990. Conserving migratory birds.
World Birdwatch 12(3)'A.
Salinas, P.J. 1980. Vertebrados poco comunes de
las zonas altas de Venezuela. Reunion
Iberoam. Conserv. Zool. Vertebr. 150pp.
Salmon, H.A. 1965. Distribution of the Jabiru in
central and northern coastal New South
Wales. Emu 65:149-151.
Salt, W.R., andJ.R. Salt. 1976. The Birds of
Alberta. Hurtig Publishers, Edmonton.
Salvadori, T. 1884. Spedizione italiana
nell'Africa aquatoriale. Risultati zoologici.
Uccelli dello Scioa e della regione fra Zeila e lo
Scioa. Ann. Mus. Civ. Genova 21:7—276.
Salvadori, T. 1899. Collezioni ornitologiche
fattenelle Isole del Capo Verde da Leonardo
Fea. Ann. Mus. Civ. Genova 30:283-312.
Salvadori, T. 1900. On the ibises of the Genus
Theristicus. Ibis (ser. 7) 6:501-517.
Salvan, J. 1967. Contribution a 1'etude des
oiseaux du Tchad. Oiseau Rev. Fr. Orn. 37:255-
284.
Salvan, J. 1971. Observationes nouvelles a
Madagascar. Alauda 39:37-42.
Salvin, O. 1865. The seabirds and waders of the
Pacific coast of Guatemala. Ibis (new ser.)
1:187-199.
Salvin, O. 1885-1886. A list of birds obtained by
Mr. Henry Whitely in British Guiana. Ibis
1885:195-219, 291-306, 418-439; 1886:57-78,
168-181,499-510.
Salzert, W. 1988. Storchenreservat im Tierpark
Rheine. Der Zoofreund 68:10-13.
Sankhala, K. 1990. Gardens of God: The Waterbird
Sanctuary at Bharatpur. Vikas Publ. House, New
Delhi. 145pp.
San Martin, P.R. 1959. Nota sobre el contenido
estomacal de un (Theristicus caudatus caudatus)
(Boddaert), 'bandurria', (Ciconiformes,
Threskiornithidae). Bio. Soc. Cienc. Nat. Taquato
l(3):79-84.
Sander, F. 1956. A list of birds of Lagos and its
environs with brief notes on their status.
Nigerian Field 21(4): 147-162.
Sanson, O., B.D. Bell, T. Andrews, and R.A.
Wilson. 1954. Visitation of Glossy Ibis.
Notornis 6:18-19.
Sapetin, Y.V. 1968. [Seasonal distribution and
migration of the Spoonbill and Glossy Ibis
according to ringing data.] Migratsii Zhivotrtykh
5:94-112.
Sarmiento, G., and M. Monasterio. 1975. A
critical consideration of the environmental
conditions associated with the occurrence of
savanna ecosystems in tropical America. Pp.
223-250 in Golley, F.B., and E. Medina (eds),
Tropical Systems. Springer-Verlag, New York.
Sasamoto, Y. 1971. A record of young Black
Stork, Ciconia nigra (Linnaeus). Tori 20:207-
208.
Sass, R.H. 1948. Glossy Ibis breeds in South
Carolina. Auk 65:459-460.
Sato, H. 1968. A note on the plumage color of the
Japanese Crested Ibis. Tori 18:301-312.
Sato, H. 1976. Toki which fly in Sado. Anima
37:22-28.
Saunders, H. 1899. Illustrated Manual of British
Birds. London.
Saunders, J.B. 1956. The Scarlet Ibis. Trinidad
Regent News G:\Q-\2.
Sauter, U., and E. Schuz. 1954.
Bestandsveranderungen beim WeiBstorch:
Dritte Ubersicht, 1939-1953. Vogelwarte 17:81-
100.
Savigny, M.J.C.L. d. 1805. Histoirie Naturelle et
Mythologique de I'Ibis. Allaca, Paris.
Savigny, M.J.C.L. d. 1810. Systeme des Oiseaux de
I'Egypte et de la Syrie. de rimprimerie
Imperriale, Paris.
Savitzky, B.P., and E.G. Samusenko (eds). 1990.
Aisty, Rasporstranenije, Ekologija, Ockrana.
Navukai Tecknika Publishing House, Minsk.
Schaeffer, P.P., and S.M. Ehlers (eds). 1980. The
Birds of Mexico: Their Biology and Conservation.
Proceedings of the National Audubon
Symposium, Tiburon, California.
Schafer, E., and W.H. Phelps. 1954. Las aves del
Parque Nacional 'Henri Pittier' (Rancho
Grande) y sus funciones ecologicas. Bol. Soc.
Venezolana Ciencias Nat. 16:3—167.
Schauffele, F., and E. Schiitz. 1968. Eine
Stelzvogel- und Scharbenkolonie im Tana-See
(Athiopien). J. Orn. 109:232-234.
Schekkerman, H., and J.CJ. van Wetten. 1987.
An ornithological winter survey of Lake Turkana,
Kenya. WIWO report 17, Nairobi.
Schell, E.H. 1968. A first Ohio record of the
White Ibis, Eudocimus albus. OhioJ. Sci. 68:17-
18.
Schenk, J. 1908. Adelekok a feher golya
369
eletmodyahoz.—Beitrage zur Lebensweise des
weissen Storches. Aquila 15:259-267.
Schenk, J. 1914. Ein Sechsergelege des Weissen
Storches. Aquila 21:269-270.
Schenk, J. 1938. Der Deutsche Storchversuch.
Proc. Int. Orn. Congr. 8:519-528.
Schenker, A. 1976. Die Waldrappkolonie im Zoo
Basel. Bull. Vereins Freunde Zool. Gartens Basel
37:9-13.
Schenker, A. 1977. Das ehemalige
Verbreitungsgebiet des Waldrapps Geronticus
eremita in Europa. Orn. Beob. 74:13-30.
Schenker, A. 1978. Hochstalter europaischer
Voegel im Zoologischen Garten Basel. Orn.
Beob. 75:96-97.
Schenker, A. 1979. Beobachtungen zur
Brutbiologie des Waldrapps (Geronticus eremita)
im Zoo Basel. Zool. Garten N. F.Jena 49:104-
116.
Schenker, A. 1981. Der Waldrapp-ein historiches
Wilbret.y^ und Hege, Naturschutz 4:7.
Schenker, A., U. Hirsch, M. Mallet, H.
Pechlaner, E. Thaler, and H. Wackernagel.
1980. Keeping and breeding the Waldrapp
Ibis. Int. Zoo News 27:9-15.
Scherf, H. 1980. Erforschungsgeschichte und
Bestandenssituation von Waldrapp und
Glattnackenrapp. Naturwiss. Rundsch. 33(2) :62-
264.
Schergalin,J. 1990. Mischbrut von Weiss-Storch
und Schwarzstorch im Zoo von Tallinn. Orn.
A/ztt. 7:177-178.
Schiefer, T.L. 1986. Wood Stork in Centre
County, Pennsylvania. Cassinia 61:76-77.
Schierer, A. 1951. Ein Massengrab von Storchen.
Ora.£*o£. 48:113-114.
Schierer, A. 1955. Ueber die kuenstliche
Aufzucht eines WeiBstorches. Vogel der Heimat
26:23-26.
Schierer, A. 1957. Les Cigognes Blanches en
Alsace de 1952 a 1956. Oiseau Rev. Fr. Orn.
27:155-160.
Schierer, A. 1958. Resultants de 10 ans de
baguage de cigognes (Ciconia ciconia) en Alsace.
Bull. Soc. Hist. Nat. Colmar 48:11-20.
Schierer, A. 1959. Efforts deployes pour la
conservation des cigognes en Alsace. Les
cigognes blanches en Alsace en 1957 et 1958.
Bull. As. Philomathique d'Alsace Lorraine suppl.
10.
Schierer, A. 1960a. Flueggerjunger WeiBstorch
draengt sich einem jungenlosen Paar auf.
Vogelwarte 20:287-288.
Schierer, A. 1960b. La forme du bee, caratere
sexuel secondaire chez la cigogne blanche
adulte (Ciconia ciconia). Oiseau Rev. Fr. Orn.
30(2): 169-172.
Schierer, A. 1960c. Beitraege aus dem Elsass zur
Kenntnis der WeiBstorch-Zugscheide.
Vogelwarte 20:288-289.
Schierer, A. 196la. Reprise d'une cigogne
blanche d'Alsace en Rhodesie du Nord. Oiseau
Rev.Fr. Orn. 31:158.
Schierer, A. 1961b. Premiers resultats des
experiences d"implantation'. en Alsace de
Cigognes blanches d'Afrique du Nord. Oiseau
Rev.Fr. Orn. 31:130-139.
Schierer, A. 1962. Quelques braves
considerations sur les Cigognes Blanches. Le
Troglodyte 1962(5):33-36.
Schierer, A. 1963. Les Cigognes blanches en
Alsace de 1959 a 1962. Alauda 31:137-148.
Schierer, A. 1967. La Cigogne Blanche (Ciconia
ciconia) en Alsace de 1948 a 1966. Lien Orn.
d'Alsace 7&8.
Schierer, A. 1972. Memoire sur la cigogne
blanche en Alsace (1948-1970). Ciconia 1:1-78.
Schierer, A. 1986. Vierzig Jahre WeiBstorch-
Forschung und -Schutz im Elsass. Beih. Veroff.
Naturschutz Landschaftspflege Bad.-Wurt. 43:329-
341.
Schierer, A. 1987. [Electric power lines and
White Storks.] Alauda 55:306.
Schierer, A. 1989a. Gigognes blanches (Ciconia
ciconia) avec caudales noires. Ciconia 13:139-
143.
Schierer, A. 1989b. Vierzig Jahre WeiBstorch-
Forschung und- Schutz im Elsass. Pp. 41-42 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status und Schutz, Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
SchifTerli, A., and A. Schierer. 1950. Die Storche
im Elsass. Orn. Beob. 47:79-93.
Schifter, H. 1984. Die Bestandsentwicklung des
Weiss-Storches, Ciconia ciconia (L.), in
Osterreich von 1975 bis 1984. Egretta 33.
Schifter, H. 1989a. Die Situation des Weiss-
Storchs in Osterreich-Stand 1988. Vogelschutz
Osterreich 4:38-39.
Schifter, H. 1989b. Zur Naturgeschichte des
Weiss-Storchs. Vogelschutz Osterreich 4:2-5.
Schiller, R.E. 1983. The Spoonbill. Zoonooz 56:
4-9.
Schlegel, H. 1863. Revue Methodique et Critique des
Collections Deposees dans cet Establissement, Vol. 5.
Museum d'Histoire Naturelle des Pays-Bas,
Leide.
Schlutheis, D. 1988. [Handrearing the Scarlet
Ibis, Eudocimus ruber.} Gefiederte Welt 112:17-18.
Schmid, B. 1931. Biologische und psychologische
Beobachtungen an Jungreihern und
Ibisvoegeln. Zool.Jahrb. Abt. Allg. Zool. Physiol.
Tiere 49:463-508.
Schmid, B. 1932. Neue biologische und
psychologische Beobachtungen an Ibisvoegeln
und Jungreihern. ZooLJahrb. Abt. Allg. Zool.
Physiol. 7^51:149-172.
Schmidt, C.R. 1968. Miscellaneous observations
on birds in Zurich Zoo, with special reference
to breeding activities. Avic. Mag. 74:61-67.
Schmidt, G.D. 1973. Resurrection of Southwellina
Witenberg, 1932, with a description of
Southwellina dimorpha sp. n. and a key to the
genera in Polymorphidae ( Acanthocephala) . J.
ParasitoL 59:299-305.
Schmidt, G.D., and A.O. Bush. 1972. Parvitaenia
ibisaesp. n. (Cestoidea: Dilepididae), from
birds in Florida./ ParasitoL 58:1095-1097.
Schmidt, K. 1989. Bigamie beim Weiss-Storch
(Ciconia ciconia) im Werratal. Orn. Mitt.
41(9):22 1-222.
Schmidt, M. 1866. Nachrichten aus dem Zool.
Garten zu Frankfurt a. m. Zeitshcrift fuer
Beobachtung. PJkge Zucht Tiere 7:421-422.
Schmidt, M. 1868. Fortpflanzung des weissen
Storches in Gefangenschaft. Zool. Garten
(Frankfurt a. M.) 9:10-23, 41-51.
Schmidt, M. 1880. On the duration of life of the
animals in the zoological garden at Frankfurt-
on-the-Main. Proc. Zool. Soc. Lond. 1880:299-
319.
Schmidt, R.A. 1980. First breeding records of the
White-faced Ibis in North Dakota. Prairie Nat.
Schmitt, B. 1967. Notes sur la biologic de la
Cigogne blanche (Ciconia ciconia) d'apres des
observations d'oiseaux captifs. Oiseau Rev. Fr.
Orn. 37:3 16-335.
Schneider, K.M. 1952. Vom Kropfstorch
(Leptoptilos, Lesson) in Gefangenschaft. Beitr.
Vogelkde 2: 196-286.
Schneider, M. 1988a. Periodisch
uberschwemmtes Dauergrunland ermoglicht
optimalen Bruterfolg des Weiss-Storches
(Ciconia ciconia) in der Save-Stromaue
(Kroatien/Jugoslawien). [Very high breeding
success of the White Stork, Ciconia ciconia, due
to temporarily flooded greenland in the
Posavina (Croatia/Yugoslavia).] Vogelwarte
34:164-173.
Schneider, M. 1988b (1989). Endangered and
rare birds in the alluvial wetlands of the Sava
River in Posavina/Croatia. Larus4Q:\67—\7S.
Schneider-Jacoby, M. and H. Ern. 1990. Saue-
Auen. Vielfalt durch Veberschwemmung. Verlag
Juergen Resch., Radolfzell, Germany.
Schnetter, W., and G. Zink. 1960. Zur Frage des
Brutreifealters Sudwestdeutschen Weiss-
Storche (Ciconia ciconia). Proc. Int. Orn. Congr.
12:662-666.
Schomburgk, R.H. 1848. Reisen in Britisch- Guiana,
Vol. 3, Versuch einer Fauna und Flora von Britisch-
Guiana. Weber, Leipzig.
Schomburgk, R. 1922-1923. Travels Through
British Guiana, 1840-1844. Daily Chronicle,
Georgetown.
Schon wetter, M. 1942. Das Ei des Balaeniceps rex
Gould. Beitr. Fortpflanzungsbiol. d. Vogel 18:41-
45.
Schonwetter, M. 1967. Handbuech der Oologie (W.
Meise, ed). Band I. Akademie-Verlag, Berlin.
Schouteden, H. 1912. Le Balaeniceps roi. Rev.
Zool. Afr. 1:347-352.
Schouteden, H. 1948-1950. De vogels van
Belgisch Congo en van Ruandra-Urundi. Les
Oiseaux du Congo Beige et du Ruanda-
Urundi. Annal. Mus. Congo Beige. Ser. 4 Zool. v.
2. Fasc 1-3.
Schouteden, H. 1954. Faune du Congo Beige et
du Ruanda-Urundi. III. Oiseaux non
Passereaux. Ann. Mus. Roy. Congo Beige,
Tervuren, Sci. Zool. 29.
Schrifter, H. 1986. Vogelparadies am Mkuzi
River/Sud Afrika. Gefiederte Welt 110:107-171.
Schrifter, H., and T. Schrifter. 1990. Die
Bestandsentwicklung des Weiss-Storches,
Ciconia ciconia (L.), in Oesterreich von 1975 bis
1984. Egretta 33:1-10.
Schroder, P. 1964. Neue Beobachtungen uber die
Begattung beim weissen Storch. Volgelkosmos
10:232-235.
Schroder, P., and G. Burmeister. 1974. Der
Schwarzs torch. Die Neue Brehm-Buecherei,
Wittenberg Lutherstadt.
Schropel, M. 1984. Aufzuchdaten fur den
WeiBstorch (Ciconia ciconia L.). Zool. Garten
54:313-329.
Schulenberg, T.S., and T.A. Parker, III. 1981.
Status and distribution of some northwest
Peruvian birds. Condor 83:209-216.
Schulz, H. 1987. Thermoregulatorisches
Beinkoten des WeiBstorchs (Ciconia ciconia).
Analyse des Verhaltens und seiner Bedeutung
fuer Verluste bei beringten Stoerchen im
afrikanischen Winterquartier. Vogelwarte
34:107-117.
Schulz, H. 1988. WeiBstorchzug: Oekologie,
Gefahrdung und Schutz des WeiBstorchs in
Afrika und Nashost. WWF Umweltforschung Ed.
3.
Schulz, H. 1989. Der Zug des WeiBstorchs-
Gefahrdung und Schutz in Afrika und Nahost.
Vogelsch. Osterreich4:1 \-\7.
Schulz, H. 1989. Zug und Ueberwinterung:
Einfuhrung. [Migration and wintering:
introduction.] Pp. 229-230 in Rheinwald, G.,
J. Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Schuster, W. 1911. Wie entleert sich der junge
Storch? Z. Orn. 21:48-49.
Schutsky, R.M. 1977. Glossy Ibis, Cattle Egret,
and Snowy Egret, three new Pennsylvania
breeding birds. Cassinia 56:3.
Schutze, B. 1984. Eine Storchengeschichte.
Panthera 1984:25-28.
Schutze, B., and J. Winkler. 1988. Erste
erfolgreiche Waldrappenaufzucht. Panthera
1988:23-27.
Schiiz, E. 1932. Wann wird der Storch
fortpflanzungsfahig? Vogelzug 3:24-29.
Schuz, E. 1934. Beobachtungen ueber
370
Paarbildung und Nestleben des Storches (C.
ciconia). Beitr. Fortpflanzungsbiol. Vogel 10:45—51.
Schiiz, E. 1935a. Beobachtungen an beringten
Stoerchen (C. ciconia) zur Brutzeit. Beitr.
Fortpflanzungsbiol. Vogel 11:61-68.
Schiiz, E. 1935b. Vernichtungen durch
Hitzewelle auf dem Zuge. Vogelzug 6:137-138.
Schiiz, E. 1936a. International Bestands-
Aufnahme am Weissen Storch 1934. Orn.
Monatsber. 44:33-41.
Schiiz, E. 1936b. The White Stork as a subject of
research. Bird-Banding 7:99-107.
Schiiz, E. 1938a. Auflassung ostpreussischer
Jungstorche in England 1936. Vogelzug 9:65-
70.
Schuz, E. 1938b. Ueber Biologic und Okologie
des Weissen Storches (C. c. ciconia). Proc. Int.
Orn. Congr. 8:577-591.
Schiiz, E. 1938c. Ueber kunstliche Verpflanzung
beiVogeln. Vogelzug 9:185-186.
Schiiz, E. 1940a. Regenwurmer als Nahrung des
Weissen Storch. Beitr. Fortpfl. 16:203-205.
Schuz, E. 1940b. Vom Zugbild des
Schwarzstorchs (Ciconia nigra). Vogelzug 11:23-
31.
Schiiz, E. 1942a. Bestandsregelnde Einflusse in
der Umwelt des Weissen Storchs (C. ciconia).
ZooLJahrb. Abt. Syst. Oekol. Geogr. 77*re 75:103-
120.
Schiiz, E. 1942b. Bewegungsnormen des Weissen
Storchs. Z. Tierpsychol. 5:1-37.
Schiiz, E. 1942c. Ueber den raumlichen und
zeitlichen Wechsel im Bestand des Weissen
Storches. [Alternation in the status of the
White Stork relating to space and time.]
Forschung. Fortschr. 18:310-312.
Schiiz, E. 1942d. Ueber die Unterscheidung
freilebender Einzelstucke des Weissen Storch.
Orn. Mber. 50:99-104.
Schiiz, E. 1943a. Geschlechtliche Ambivalenz
beim Weissen Storch. Vogelzug 14:134-137.
Schiiz, E. 1943b. Storche als Gesundheitspolizei.
Vogelwelt 68:90-91.
Schiiz, E. 1943c. Ueber die Jungenaufzucht des
Weissen Storches (C. ciconia). Z. Morphol.
Oekol. Tiere 40:181-237.
Schiiz, E. 1943d. Versuche ueber die Bindung
des Storch an seinen Aufzuchtort. Vogelzug
14:137-141.
Schiiz, E. 1943e. Vierzehn Jahre Storchsiedlung
Rossitten. Vogelzug 14:90-109.
Schiiz, E. 1944. Nesterwerb und Nestbesitz beim
Weissen Storche. Z. Tierpsychol. 6:1-25.
Schiiz, E. 1948. Stoerche als Eingeborene-Beute.
Vogelwarte 15:8-18.
Schuz, E. 1949a. Die Spaet-Auflassung
ostpreussischer Jungstoerche in West-
Deutschland durch die Vogelwarte Rossitten
1933. Vogelwarte 15:63-78.
Schuz, E. 1949b. Hilfe fur Freund Adebar.
Merkblaeter f. Angew. Vogelkunde 6:1-6.
Schiiz, E. 1949c. Reifung, Ansiedlung und
Bestandswechsel beim Weissen Storch (C.
ciconia). Pp. 217-228 in Mayr, E., and E.
Schiiz (eds), Ornithologie als Biologische
Weissenschaft. Heidelberg.
Schiiz, E. 1950. Die Fruhauflassung
ostpreussischer Jungstorche in West-
Deutschland durch die Vogelwarte Rossitten
1933-1936. Bonn. Zool. Beitr. 1:239-253.
Schiiz, E. 195la. Dreizehn Ringfunde von Weiss-
Storchen. Vogelwarte 16:79-82.
Schuz, E. 1951b. Uberblick uber die
Orientierungsversuche der Vogelwarte
Rossitten (jetzt: Vogelwarte Radolfzell). Proc.
Int. Orn. Congr. 10:249-268.
Schiiz, E. 1951c. Vogel als Traeger von
Eingeborenen-Pfeilen. Vogelwarte 16:77-79.
Schuz, E. 1952. Zur Methode der
Storchforschung. Beitr. Vogelkde 2:287-298.
Schiiz, E. 1953a. Die Zugscheide des Weissen
 Storches nach den Beringungs-Ergebnissen.
Bonn. Zool. Beitr. 4:31-72.
Schiiz, E. 1953b. Gestorte Jungenaufzucht beim
Weissen Storch in Rosswag.yA. Ver. VaterL
Naturk. 108:120-125.
Schiiz, E. 1954a. Jungstoerche als Opfer
schwarmender Bienen. Vogelwarte 17:217-218.
Schiiz, E. 1954b. Schadigt der Ausfall des
Chamsina den Heimzung des WeiBstorchs?
Vogelwarte 17:166-168.
Schiiz, E. 1954c. Vor dem Ende des Storches
(Ciconia c. bqyciand) Japan? Vogelwarte 17:164.
Schuz, E. 1954d. Zur Verbreitung des Weiss-
Storchs im Oberrheingebiet. Jh. Ver, VaterL
Naturk. 109:141-144.
Schiiz, E. 1955a. Ueber den Alteraufbau von
WeiBstorchpopulation en. Proc. Int. Orn. Congr.
11:522-528.
Schuz, E. 1955b. Vom Zug des WeiBstorchs im
Raum Syrien bis Agypten. Vogelwarte 18: 5-13.
Schiiz, E. 1956. Stoerche am Federsee in Alter
Zeit. Landess. Naturschutz Landschaftspfl. 24:257-
264.
Schiiz, E. 1957a. Das Verschlingen eigener
Junger ("Kronismus") bei Vogeln und seine
Bedeutung. Vogelwarte 19:1-15.
Schiiz, E. 1957b. Vogelkunde am Manyas—See
(Turkei). Vogelwarte 19:41-44.
Schuz, E. 1959a. Problems about the White
Stork, C. ciconia, in Africa seen from an
European view point. Proc. 1957 Pan-Afr. Orn.
Congr. (= Ostrich suppl. 3) 1:333-341
(Livingstone, Cape Town 1957).
Schiiz, E. 1959b. Stoerche und Lokustizide.
Vogelwarte 20:182-184.
Schuz, E. 1960. Die Verteilung des WeiBstorchs
im sudafrikanischen Ruheziel. Vogelwarte
20:205-222.
Schuz, E. 1962. Ueber die nordwestliche
Zugscheide des Weissen Storchs. Vogelwarte
21:269-290.
Schiiz, E. 1963a. On the northwestern migration
divide of the White Stork. Proc. Int. Orn. Congr.
13:475-480.
Schuz, E. 1963b. Ueber die Zugscheiden des
WeiBstorchs in Afrika, Ukraineund Asien.
Vogelwarte 22:65-67.
Schiiz, E. 1963c. Zum zweitenmal ein
Brutvorkommen des WeiBstorchs in Sudafrika.
Vogelwarte 22:38-39.
Schiiz, E. 1964. Zur Deutung der Zugscheiden
des Weiss-Storchs. Vogelwarte 22:194-223.
Schiiz, E. 1965a. Gefahren fur den ziehenden
WeiBstorch. Int. Rat F. Vogelschutz, Dtsche Sek.
5:15-22.
Schiiz, E. 1965b. Von 'Oshima' vom
Schwarzschnabelstorch und von der
Lebensdauer der Storche. Beitr. Vogelkde
10:329-333.
Schiiz, E. 1966. Ueber Palaarkten in Athiopien
im April, mit zusatzlichen Angaben uber den
WeiBstorch. Vogelwarte 23:285-289.
Schiiz, E. 1967a. Ornithologischer April-Besuch
in Athiopien, besonders am Tanasee. Stuttg.
Beitr. Naturkd. 171:1-22.
Schiiz, E. 1967b. Verbreitungsgrenzen der
Westrasse des Weiss-Storchs (C. c. ciconia).
Vogelwarte 24:116-122.
Schuz, E. 1968a. Der Zug des Weiss-Storchs.
Schulwandbild in der Reihe. Der Neue
Schulmann, Nr. 4184, dazu Textheft, Lieferung
43:9-16. Verlag Holzwarth, Stuttgart.
Schiiz, E. 1968b. Eine neue Vogelkolonie am
Tanasee in Athiopien. Kosmos 64:368-369.
Schiiz, E. 1968c. Ornithologischer Oktober-
Besuch am Tanasee (undin bei addis Ababa),
Aethiopien. Stuttg. Beitr. Naturkd. 189:1-43.
Schiiz, E. 1969. Storks and other birds carrying
arrowheads. Ostrich 40:17-19.
Schuz, E. 1970a. The riddle of the so-called
'Benin Ibis' and the artificial wattled birds.
Ostrich, suppl. 8:15-19.
Schuz, E. 1970b. Glossy Ibis and Hermit Ibis
(Geronticus eremita) were found in Egypt. Proc.
Ill Pan-Afr. Orn. Conf. Ostrich 8.
Schiiz, E. 1971. Grundriss der Vogelzugskunde. Paul
Parey, Berlin. 390pp.
Schiiz, E. 1973. White Stork colonization—a
social factor also? Bokmakierie 25:69-70.
Schiiz, E. 1975. Greifvogel als Angreifer von
Weifistorchen (Ciconia ciconia). Vogelwelt
96:114-116.
Schuz, E. 1977. Changes in the status of the
White Stork. ICBP President's Letter 40:2-3.
Schuz, E. 1979a. Rettet den WeiBstorch.
Naturfors. Gesell. Rheinaubund, Schaffhausen,
Fulgblatt-Ser. 7/15:1-44.
Schiiz, E. 1979b. Results of the III international
census (1974) of the White Stork. Bull. ICBP
13:173-179.
Schiiz, E. 1980. Status un Veranderung des
WeiBstorch-Bestandes. Naturwiss. Rundsch.
33:102-105.
Schuz, E. 1981a. Noch ein Kapitel WeiBstorch.
Pp. 77-91 in Kahl, M.P., Welt der Storche.
Verlag P. Parey, Hamburg.
Schuz, E. 1981b. The protection of the White
Stork in African countries. Okol. Vogel 3:307-
310.
Schiiz, E. 1983. (Review of) Thiede, Ulrike
(1982): Japan-Ibis und Japanische Nachtigall
als Beispiele zweir Pole im Naturverstandnis
derjapaner. J. Orn. 124:96.
Schiiz, E. 1984a. Schwarzschnabelstorch Ciconia
boyciana, 2. Bericht. Vogelwarte 32(4):3\7-3\9.
Schiiz, E. 1984b. Ueber Syngenophagie,
besonders Kronismus: ein Beitrag zur
Ethologie speziell des WeiBstorchs. Okol. Vogel
6:141-158.
Schiiz, E. 1985a. Ueber die sogenannten
Storchgerichte. Orn. Mitt. 37:352-357.
Schiiz, E. 1985b. Vom Schwarzschnabelstorch
des Fernen Ostens. Gefoderte Welt 109:32.
Schiiz, E. 1986a. Anmerkungen zu: Gibt es eine
Bigamie beim Weiss-Storch (Ciconia ciconia)?
Orn. Mitt. 38:139.
Schiiz, E. 1986b. Buecher ueber den WeiBstorch
(Ciconia ciconia). Beih. Verojf. Naturschutz
Landschaftspfiege Bad.-Wurtt. 43:351-360.
Schiiz, E. 1986c. Danemark: Einst das klassische
Storchland. Beih. Verojf. Naturschutz
Landschaftspjkge Bad.-Wurtt. 43:127-130.
Schiiz, E. 1986d. Ueber die Namen des WeiB-
storchs (Ciconia ciconia). Beih. Veroff. Naturschutz
Landschaftspjkge Bad.-Wurtt. 43:15-24.
Schiiz, E. 1986e. Zur bibiographie des Weiss-
storchs (C. ciconia). Beih. Veroff. Naturschutz
Landschaftspjkge Bad.-Wurtt 43:361-372.
Schuz, E., and P. Berthold. 1966. Ueber das
Bruten Fernziehender Vogelarten an beiden
Polen der Jahresverbreitung. Vogelwarte
23:233-234.
Schuz, E., and R. Bohringer. 1950. Vom Zug des
Weiss-Storchs on Afrika und Asien nach den
Ringfunden bis 1949> Vogelwarte 15:160-187.
Schiiz, E., and W. Gehlhoff. 1967. Die
Brutverbreitung des WeiBstorchs im Vorderen
und Mitteren Orient. Vogelwarte 24:48-63.
Schiiz, E., and T. Schiiz. 1932a. Von der
Ankunft des Weissen Storch am Brutplatz.
Orn. Beob. 29:117-1^0.
Schuz, E., and T. Schiiz. 1932b.
Brutbeobachtungen am Storch (Ciconia ciconia).
Beitr. Fortpflanzungsbiol. Vogel 8:18-21.
Schiiz, E., and J. Szijj. 1960a. Vorlaufiger
Bericht ueber die Internationale
Storchbestandsaufnahme des WeiBstorchs
1958. Vogelwarte 20:253-257.
Schuz, E., and J. Szijj. 1960b.
Bestandsveranderungen beim WeiBstorch:
Vierte Uebersicht, 1954-1958. Vogelwarte
20:258-273.
Schuz, E., and J. Szijj. 1961. Vom WeiB-
371
storchbestand in Deutschland 1934 bis 1958. J.
Orn. 102:28-33.
Schiiz, E., and J. Szijj. 1962. Report on the
international census of the White Stork 1958.
Bull. Int. Comm. Bird Preserv. 8:86-98.
Schuz, E., and J. Szijj. 1975.
Bestandsveranderungen beim WeiBstorch
Funfte Uebersicht: 1959-1972. Vogelwarte
28:61-93.
Schuz, E., and H. Veroman. 1982.
Mittelasiatischer WeiBstorch. Vogelwarte
31:467-469.
Schuz, E., and G. Zink. 1955. Verzeichiswerk
1955. C. Bibliographic der WeiBstorch-
Untersuchungen der Vogelwarten Rossitten-
Radolfzell und Helgoland. Vogelwarte 18:81^85.
Schuz, E., H. Lange, W. Gaupp, and H. Sick.
1932. Brutnachweise dreijahriger Stoerche.
Vogelzug3:\76-l77.
Schiiz, E., F. Burr, H. Lange, H. Johansen, and
R. Kuhk. 1955. Hohe Lebensalter bei
Storchen. Vogelwarte 18:21-22.
Schiiz, E.5 M. Casement, and H. Seilkopf. 1963.
Weisser Storch: Falle von Suesgolf-Querung
und Sinai-Zug. Vogelwarte 22:26-30.
Schwartz, A., and R.F. Kinikowski. 1963.
Observations on West Indian birds. Proc. Acad.
Nat. Sci. Phila. 115(3).
Schwitulla, H. 1962. Heuschreckenbekampfung
und Stoerche. Gesunde Pflanzen 14:70-71.
Sciple, G.W. 1963. Changing status of
occurrence of the Glossy Ibis. Oriole 28:23-25.
Sclater, P.L. 1857. On a collection of birds
received by M. Salle from southern Mexico.
Proc. Zool. Soc. Lond. 1857:226-237.
Sclater, P.L. (ed.). 1867. Nitzsch's Pterylogy. Ray
Society, London.
Sclater, P.L. 1869. On the breeding of birds in
the gardens of the Zoological Society of
London during the past twenty years. Proc.
Zool. Soc. Lond. 1869:626-629.
Sclater, P.L. 1878. Notes on the breeding of the
Sacred Ibis in the Zoological Society's
gardens. Ibis 1878:449-451.
Sclater, P.L. 1906. On some birds collected by
Mr. Douglas Carruthers in the Syrian Desert.
Ibis 6:307-317.
Sclater, P.L. 1912. On the birds collected by Mr.
Claude H.B. Grant at various localities in
South Africa-Part IV. Ibis 9:1-63.
Sclater, P.L., and W.H. Hudson. 1889. Argentine
Ornithology, vol. 2. R.H. Porter, London.
251PP.
Sclater, W.L. 1896. A visit to Dassen Island, the
home of the Jackass Penguin. Ibis (ser. 7)
2:519-525.
Sclater, W.L. 1930. Systema avium
Aethiopicarum. A systematic list of the birds of
the Ethiopian Region. Part 1. Taylor and
Francis, London.
Sclater, W.L., and C. Mackworth-Praed. 1920. A
list of the birds of the Anglo-Egyptian Sudan,
based on the collections of Mr. A.L. Butler,
Mr. A. Chapman and Capt. H. Lynes, R.N.,
and Major Cuthbert Christy, R.A.M.C. (T.F.)
Part IV. Concluded. Pelecanidae-
Struhionidae. Ibis 11:781-855.
Sclater, W.L., and R.E. Moreau. 1932.
Taxonomic and field notes on some birds of
the north-eastern Tanganyika Territory. Ibis
(ser. 13) 2:487-522.
Sclater, W.L., and R.E. Moreau. 1933a. On
some birds of Northwestern Tanganyika
Territory. Ibis 2:420-421.
Sclater, W.L., and R.E. Moreau. 1933b.
Taxonomic and field notes on some birds of
north-eastern Tanganyika Territory. Part V.
Ibis (ser. 13) 3:399-440.
Scott. D.A. 1980. A Preliminary Inventory of
Wetlands of International Importance for Waterfowl
in West Europe and Northwest Africa. IWRB
 Spec. Publ. No. 2. Slimbridge, Glos, England.
127pp.
Scott, D.A. 1982. Wetlands and waterfowl in
South America: a continental perspective.
Unpublished report, IWRB Symposium
Edmonton, Canada, 25-28 May, 1982.
International Waterfowl Research Bureau,
Slimbridge.
Scott, D.A. (ed.). 1989. A Directory of Asian
Wetlands. IUCN, The world Conservation
Union, Gland, Switzerland.
Scott, D.A., and M. Carbonell. 1986. A Directory
of Neotropical Wetlands. IUCN, Cambridge,
United Kingdom.
Scott, D.A., and C.M. Poole. 1989. A Status
Overview of Asian Wetlands. Asian Wetland
Bureau Publication No. 53. Kuala Lumpur,
Malaysia.
Scott, D.A., and P.M. Rose. 1989. Asian waterfowl
census 1989, mid-winter waterfowl counts in southern
and eastern Asia January 1989. IWRB,
Slimbridge, U.K.
Scott, J. A. 1972. Woolly-necked Stork breeding
in Rhodesia. Ostrich 43:68.
Scott, J.A. 1975. Observations on the breeding of
the Woollynecked Stork. Ostrich 46:201-207 .
Scott, S.L. 1987. Field Guide to the Birds of North
America. National Geographic Society,
Washington, D.C.
Scott, W.E.D. 1887. The present condition of
some of the bird rookeries of the gulf coast of
Florida. Auk 4:135-144.
Scott, W.E.D. 1889. A summary of observations
on the birds of the gulf coast of Florida. Auk
6:13-18.
Scott, W.E.D., and Sharpe, R.B. 1912. Pp. 345-
504. in Reports of the Princeton University
Expeditions to Patagonia, 1896-1899. II.
Ornithology Part III. Charadriidae—Anatidae. The
University of Princeton, New Jersey.
Seebohm, H. 1886. On the identity of Ibis
propinqua with Ibis melanocephala. Ibis 6(24):473-
474.
Seebohm, H. 1890. The Birds of the Japanese
Empire. R.H. Porter, London. 386pp.
Sefton, A.R. 1958. Bird notes from the Illawarra
District, New South Wales. Emu 58:393-394.
Sefton, A.R., and J.A. Devitt. 1962. Additions to
the birds recorded from the Illawarra District.
£WM 62: 184-187.
Seibt, U., and W. Wickler. 1978. Marabou storks
wash dung beetles. Z. Tierpsychol. 46(3):324-
327.
Seitz, A. 1940. Der Hausstorchbestand in
Burgenland (Gau Niederdonau) 1934-1939.
Zeit. Beitr. Fortpflanzungsbiol. Vogel 16:87-92.
Sell, R.A. 1917. Some notes on the effects upon
bird life, of the Corpus Christi storm of August
18, 1916. Condor 19:43-46.
Sell, R.A. 1918. The Scarlet Ibis in Texas. Condor
20:78-82.
Semple, J.B. 1932. Glossy Ibis in Florida. Auk
49:337-339.
Serle, W. 1943. Further field observations on
northern Nigerian birds. Ibis 85:264-300.
Serle, W. 1965. A third contribution to the
ornithology of the British Cameroons. Ibis
107:60-94.
Serle, W., and G.J. Morel. 1984. A Field Guide to
the Birds of West Africa. Collins, London.
Serle, W., G.J. Morel, and W. Hartwid. 1977. A
Field Guide to the Birds of West Africa. Collins,
London.
Serna D., 1984. Avifauna Parcial de la Guajira.
Museo de Historia Natural, Medellin,
Colombia. 52pp.
Serna D., 1990. Algunas aves observadas en la
Cienaga de Ayapel, Cordoba. Bol. S.A.O.
Serrao, J.S. 1986. The White Stork Ciconia ciconia.
Serventy, D.L., and H.M. Whittell. 1962. Birds of
Western Australia, 3rd edn. Paterson Brokensha,
Perth. 427pp.
Serventy, V. 1958. Bird notes from the
Dumbleyung Camp-out, 1956. Emu 58:5-20.
Seward, D. 1987. Europe's vanishing storks. Int.
Wildl. 17(3):4~11.
Seys, G., and E. Lincent. 1932. La collection
d'oiseaux du Musee Hoangho Paiho de Tien-
Tsin. Mission de sein Hsein.
Shackford, J.S. 1991. The Roseate Spoonbill in
Oklahoma. Bull. Okla. Ornithol. Soc. 24:1-3.
Shah, R.V., and J.H. Desai. 1972 (1975).
Possible explanation for voicelessness in the
adult stork, Ibis leucocephalus. Pavo 10:21-29.
Shah, R.V., and J.H. Desai. 1975a (1976).
Growth and development of the Painted Stork,
Ibis leucocephalus Pennant. I. Embryonic
development. Pavo 13(2):83-87.
Shah, R.V., and J.H. Desai. 1975b (1976).
Growth and development of Painted Stork Ibis
leucocephalus Pennant. II. Post hatching growth
pattern and motor development. Pavo
13(2):88-101.
Shah, R.V., G.K. Menon, J.H. Desai, and M.B.
Jani. 1977. Feather loss from capital tracts of
Painted Storks related to growth and maturity.
Part 1. Histo-physiological changes and lipoid
secretion in the integument. J. Anim. Morphol.
Physiol. 24:98-107.
Shanholtzer, F.G. 1970. Breeding records and
distribution of the Glossy Ibis on the Georgia
coast. Oriole 35:37-39.
Shanholtzer, F.G., WJ. Kuenzel, and JJ.
Mahoney. 1970. Twenty-one years of the
McKinney's Pond Rookery. Oriole 35:23-28.
Shannon, P.W. 1987. The Jabiru Stork (Jabiru
mycteria) in zoo collections in the United States.
Colonial Waterbirds 10:242-250.
Shannon, P. 1988. Who delivers the storks?
Jabiru Stork management at the Audubon
Zoo. At the Zoo Summer 1988.
Sharland, M. 1957a. White Ibis in Tasmania.
Emu 57:301-302.
Sharland, M. 1957b. Royal Spoonbill in
Tasmania. Emu 57:234-235.
Sharland, M. 1957c. Egrets at Ulmarra, N.S.W.
Emu 57:295-301.
Sharland, M. 1958. Branch Reports. Emu 58:
43-44.
Sharma, P.N. 1980. Host-parasite relations of
Chaunocephalus similiferox, Verma 1936
(Trematode: Echinostomatidae) in the bird
host Anastomus oscitans. Acta Parasitol. Polonica
27(29-^5):273-278.
Sharpe, R.B. 1895. On a collection of birds made
by Dr. A. Donaldson Smith during his recent
expedition in Western Somaliland. Proc. Zool.
Soc. Lond. 1895:457-520.
Sharpe, R.B. 1898. Catalogue of the Birds in the
British Museum, vol. 26. British Museum
(Natural History), London. 687pp.
Sharpe, R.B. 1899. A Hand-list of the Genera and
Species of Birds, vol. 1. British Museum
(Natural History), London.
Sharrock, E.M. 1982. Observation of a Glossy
Ibis Plegadisfalcinellus. Alauda 50:70.
Shaw, G. 1800. Description of a new species of
Mycteria [senegalensis]. Trans. Linn. Soc. Lond.
5:32-35.
Shaw, T.H. 1936. The Birds ofHopei Province. Fan
Memorial Inst. Biology, Peking.
Sheldon, F.H. 1987. Phylogeny of herons
estimated from DNA-DNA hybridization
data. Auk 104:97-108.
Shelley, G.E. 1871. Contribution to the
ornithology of Egypt. Ibis (ser. 3) 1:38-54,
131-147,309-319.
Shelley, G.E. 1872. A Handbook of the Birds of
Egypt. John Van Voorest, Paternoster Row,
London.
Shepard, J.B. 1962. Check list of birds of
Swartberg District. S. Afr. Avifauna ser. 6.
372
Sheshukumar, Y. 1984. Feeding ecology of the
Indian Black Ibis in and around Saurashtra
University Campus. M.S. Dissertation.
Saurashtra University, Rajkot, India.
Shi, Z. 1991. Preliminary observation on the
winter foraging ecology of Oriental White
Storks (Ciconia bqyciana) in Jiangsu Province,
China. Pp. 107-114, in Coulter, M.C., Q.
Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork
(Ciconia bqyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Shibaev, I.V., and N.M. Litvenenko. 1977. [New
information on the spreading of Japanese
Cranes (Grusjaponica) and Oriental White
Storks (Ciconia bqyciana).] Pp. 251-252 in
Winter, S. (ed.), VII All-Soviet Ornithological
Conference. Zoological Institute of the Academy
of Science of the Ukranian SSR, Kiev.
Shibaev, I.V., N.N. Semenchenko, and V.A.
Limin. 1976. [Nesting area of the White Stork
on the right bank of the Ussuri River.] In
Nature Conservation in the Soviet Far East.
Shibata, T. 1969. Declaration of Emergency—
Lets' save Toki. Nature (Jap. League Prot. Birds)
208:15.
Shichiri, S. 1980. Handraising the Black-headed
Ibis. Animals Zoos 32:150-151.
Shields, M.A. 1985. An analysis of Fish Crow
predation on eggs of the White Ibis at Battery
Island, North Carolina. M.S. Thesis.
University of North Carolina, Chapel Hill,
North Carolina.
Shields, M.A. 1987. Internest displacement of
White Ibis eggs. Wilson Bull. 99:273-275.
Shields, M.A., and R.D. Bjork. 1986. The
breeding birds of Battery Island, N.C.: an
update. Chat 50:101-104.
Shields, M.A., and J.F. Parnell. 1983. Expansion
of White Ibis nesting in North Carolina. Chat
47:101-103.
Shields, M.A., and J.F. Parnell. 1986. Fish Crow
predation on eggs of the White Ibis at Battery
Island, North Carolina. Auk 103:531-539.
Short, L.L., Jr. 1966. A new Pliocene stork from
Nebraska. Smithson. Misc. Collect. 149:1-11.
Short, L.L. 1975. A zoogeographic analysis of the
South American Chaco avifauna. Bull. Am.
Mus. Nat. Hist. 154.
Short, L.L., J.F.M. Home, and C. Muringo-
Gichuki. 1990. Annotated check-list of the
birds of East Africa. Proc. West. Found. Vertebr.
Zool. 4:61-246.
Shortridge, G.C. 1905. Birds collected around
Hanover, Cape Colony, from July 20th to the
end of September 1903.J. S. Afr. Orn. Union
2:55-83.
Shufeldt, R.W. 1900. Osteology of the
Herodiones. Ann. Carnegie Mus. 1:158-249.
Shufeldt, R.W. 1901. Notes on the osteology of
Scopus umbretta and Balaeniceps rex. f. Anat.
Physiol. 2:405-412.
Sibley, C.G. 1960a. The taxonomic relationships
of birds as indicated by the egg-white proteins:
a request for help. Avic. Mag. 1960:52-56.
Sibley, C.G. 1960b. The electrophoretic pattern
of avian egg-white proteins as taxonomic
characters. Ibis 102:215-284.
Sibley, C.G. 1989. Response to E. Mayr. Auk
106:512-515.
Sibley, C.G., and J.E. Ahlquist. 1972. A
comparative study of the egg white proteins of
non-passerine birds. Peabody Mus. Nat. Hist.
Yale Univ. Bull. 39:1-276.
Sibley, C.G. and J.E. Ahlquist. 1983. The
phylogeny and classification of birds based on
the data of DNA-DNA hybridization. Pp.
245-292 in Johnston, R.F. (ed.), Current
Ornithology, vol. 1. Plenum Press, New York.
Sibley, C.G., and J.E. Ahlquist. 1985. The
relationships of some groups of African birds,
based on comparisons of the genetic material,
 DNA. Pp. 115-161 in Schuchmann, K.-L.
(ed.), African Vertebrates: Systematics, Phytogeny,
and Evolutionary Biology. Zool.
Forschungsinstitut und Museum A. Koenig,
Bonn.
Sibley, C.G., and J.E. Ahlquist. 1990. Phytogeny
and Classification of Birds. Yale University Press,
New Haven, Connecticut. 1008pp.
Sibley, C.G., and B..L. Monroe, Jr. 1990.
Distribution and Taxonomy of Birds of the World.
Yale University Press. New Haven,
Connecticut. 1136pp.
Sibley, C.G., K.W. Corbin, and J.H. Haavie.
1969. The relationships of the flamingos as
indicated by the egg-white proteins and
hemoglobins. Condor 71:155-179.
Sibley, C.G., J.E. Ahlquist, and B.L. Monroe, Jr.
1988. A classification of the living birds of the
world based on DNA-DNA hybridization
studies. Auk 105:409-423.
Sick, H. 1967. Hochwasserbedingte
Vogelwanderungen in den neuweltlichen
Tropen. Vogelwarte 24:1-6.
Sick, H. 1968. Vogelwanderungen im
kontinentalen Sudamerika. Vogelwarte 24:217-
243.
Sick, H. 1969. Aves brasileiras ameacadas de
extincao e nocoes gerais de conservacao de
aves do Brasil. An. Acad. Bras. Cienc. 41:205-
229.
Sick, H. 1972. A ameaca da avifauna brasileira.
Pp. 99-153 in Especies da Fauna Brasileira
Ameacadas de Extincao. Academia Brasileira de
Ciencias, Rio de Janeiro.
Sick, H. 1985. Ornitologia Brasileira, Uma
Introducao. Editora Universidade de Brasilia,
Brasilia.
Sick, H., and D.M. Teixeira. 1979. Notas sobre
aves brasileiras raras ou ameacadas de
extincao. Publ. Avulsas Mus. Nac. (Rio dej.)
62:1-39.
Siebold, P.P. 1850. Fauna Japonia Vol. 5, Oiseaux de
Japan, Tokyo.
Siefke, A. 1974. Zweiter Fund eines europaischen
Weifistorches in Indien. 7^/^21:139.
Siefke, A. 1981. Dispersal and site tenacity of the
White Stork Ciconia ciconia according to ringing
results from East Germany. Zoot.Jahrb. Abt.
Syst. Okol. Geogr. Tiere 108:15-35.
Siegfried, W.R. 1966a. The present and past
distribution of the Bald Ibis in the Province of
the Cape of Good Hope. Ostrich 37:216-218.
Siegfried, W.R. 1966b. The Bald Ibis. Bokmakierie
18:54-57.
Siegfried, W.R. 1967. The distribution and status
of the Black Stork in Southern Africa. Ostrich
38:179-185.
Siegfried, W.R. 1968. Spoonbills breeding at
Stellenbosch, Cape Province, South Africa.
Ostrich 39:\99.
Siegfried, W.R. 1971. The status of the Bald Ibis
in southern Africa. Biol. Conserv. 3:88-91.
Siegfried, W.R. 1972. Discrete breeding and
wintering areas of the Waldrapp Geronticus
eremita (L.). Bull. Br. Orn. Club 92:102-103.
Siegfried, W.R., P.G.H. Frost, J. Cooper, and
A.C. Kemp. 1976. South African Red Data
Book—Aves. South African National Science
Program Report, No. 7, 108pp.
Siewert, H. 1932a. Beobachtungen am Horst des
Schwarzen Storches. J. Orn. 80:533-541.
Siewert, H. 1932b. Stoerche. D. Reimer/E.
Vohsen, Berlin.
Siewert, H. 1955. Stoerche. Erlebnisse mil dem
Schwarzen und Weissen Storch. C. Bertelsmann,
Gutersloh.
Silva-Porto, F.C., and D.M. Teixeira. 1984. Urn
estudo comparative preliminar sobre as
avifaunas das restingas do leste do Brasil. Pp.
343-349 in Lacerda et al. (eds), Restingas:
Origem, Estruturas, Processes. CEUFF, Niteroi,
Rio de Janeiro.
Silvestri, A. 1977. Nidificazione de cigogne
ambientate in Romagna. [Nesting of the white
stork in Romagna.] Riv. Ital Orn. 47(3):250-
253.
Silvius, MJ. 1985. West Peninsular Malaysia.
Interwader Newsletter 5:6.
Silvius, MJ. 1985. South-east Sumatra.
Interwader Newsletter 5:6.
Silvius, MJ. 1986. Survey of the coastal wetlands
in Sumatra Selatan and Jambi, Indonesia.
PHPA-Interwader Report No. 1.
Silvius, M J. 1988. On the importance of
Sumatra's east coast for waterbirds. Kukila
3:117-137.
Silvius, M J., and W J.M. Verheugt. 1985.
Mangroven. Natuur Tech. 53:576-587.
Silvius, M J., and W J.M. Verheugt. 1986. The
birds of the Berbak Game Reserve, Jambi
Province, Sumatra. Kukila 2:76-84.
Silvius, M J., and WJ.M. Verheugt. 1990. The
status of storks, ibises and spoonbills in
Indonesia. Kukila 4:119-132.
Silvius, MJ., WJ.M. Verheugt, and J. Iskandar.
1985. Coastal surveys in southeast Sumatra.
Interwader Annu. Rep. 1984:133-142.
Silvius, M J., WJ.M. Verheugt, and J. Iskandar.
1986. Coastal wetlands inventory of southeast
Sumatra. ICBP Study Rep. 9.
Simeonov, S.D., and S.P. Dimitrova. 1987. A
study of the number of White Storks (Ciconia
ciconia (L.)) in the district of Pernik. Ekologiya
(Sofia) 19:26-34.
Simmons, F.B. 1948. A list of birds observed in
Chittagong, E. Bengal, during 1944 and 1945.
J. Bombay Nat. Hist. Soc. 47:637-644.
Simon, A.L. 1952. A Concise Encyclopedia of
Gastronomy. Harcourt, Brace and Co., New
York.
Simon, N. and F. Geroudet. 1970. Last Survivors.
World Publ., New York.
Simpson, M.B. Jr. 1966. Status of the Scarlet
Ibis in South Carolina: historical records from
John Abbot and Alexander Wilson. Chat 52:4-
5.
Simpson, M.B., and Dat. 1984. The Birds of
Australia. Lloyd O'Neil PTy, Victoria,
Australia.
Singh, G., and C. Singh. 1960. The Adjutant
Stork, Leptoptilos dubius (Gmelin), a destroyer of
locusts in Raj as than. J. Bombay Nat. Hist. Soc.
57:221-222.
Singleton, R.T. 1973. Species composition and
behavior of herons and egrets at the Burdette
Heronry in northeastern Arkansas. M.S.
Thesis, Arkansas State University, Arkansas,
USA.
Sitwell, N. 1984. Storks get a helping hand. Int.
Wildl. 14:14-19.
Skaar, P.D. 1975. Montana Bird Distribution. P.D.
Skaar, Bozeman, Montana.
Skadsen, D. 1988. White-faced Ibis nesting in
Day County, South Dakota. Bird Notes 40:94.
Skead, C J. 1951. A study of the Hadedah Ibis
Hagedashia h. hagedash. Ibis 93:360-382.
Skead, C J. 1954. African Spoonbills (Platalea
alba) nesting near Graaf Reinet. Ostrich 25:41.
Skead, C J. 1966. Hadedah Ibis Hagedashia
hagedash (Latham) in the eastern Cape
Province. Ostrich 37:103-108.
Skead, C J. 1967. Ecology of birds in the eastern
Cape Province. Ostrich suppl. 7:1-103.
Skinner, M.P. 1926. Wood Ibis in the
Yellowstone National Park. Condor 28:99.
Skokova, N.N. 1959. [The Spoonbill in the Volga
Delta, its ecology and a role in the fishery.]
Ornitologia 2:262-270.
Skokova, N.N. 1963. On the quantitative study
of feeding in piscivorous birds. Ornitologiya
4:288-296.
Skoptsov, V. 1987. [The storks (Ciconia spp.) in
Tambov Region.] Ornitologiya 22:194-195.
Skov, H. 1979. Red storken i Danmark
373
Feltornitologen. Panurus (Dansk Orn.
Skov, H. 1981. Storken i Sonderjylland. Panurus
(Dansk Orn. Foren.) 1:9-28.
Skov, H. 1982. Opsaetning af reder kan nytte.
Dansk Orn. Foren., Fugle 2:12-13.
Skov, H. 1983. Storkens status i Sdr. jylland
1983. Panurus (Dansk Orn. Foren.) 3:8-10.
Skov, H. 1984. Regional reports about White
Storks (Ciconia ciconia)— Denmark. World
Working Group on Storks, Ibises and
Spoonbills. ICBP Report 2:8.
Skov, H. 1985. Storkens status i Sdr. jylland
1984. Panurus (Dansk Orn. Foren.) 1:18-21.
Skov, H. 1986a. Storkens status i Sdr. jylland
1985. Panurus (Dansk Orn. Foren.) 2:13-16.
Skov, H. 1986b. Storken i Sonderjylland.
Sonderjyllands Fugleliv. Forlaget Fulica,
Aabenraa:\27-\36.
Skov, H. 1986c. Storken sidder ikke pa den
moderne landmands tag. Dansk Orn. Foren.
Fugle 3:4-5.
Skov, H. 1987a. Storkens status i Sonderjylland i
1987. Panurus (Dansk Orn. Foren.) 3:12-14.
Skov, H. 1987b. Storke og eldoden. Dansk Orn.
Foren., Fugle 3:16.
Skov, H. 1987c. Sidste udkald for de nordjyske
storke. Riden (Dansk Orn. Foren. Nordjyllands
Ami) 4:10-13.
Skov, H. 1987d. Det sydlige Afrika— noglen til
Storkens fremtid? Dansk Orn. Foren., Fugle
1:30-31.
Skov, H. 1987e. Vinterstorke. Hjejlen (Dansk Orn.
Foren. Vigorg Amt) 1:9-11.
Skov, H. 1987f. Storkereder. Dansk Orn. Foren.,
Fugle 5:12-14.
Skov, H. 1988a. Storkens status i Syd- og
Sonderjylland i 1988. Panurus (Dansk Orn.
Foren.) 3:17-20.
Skov, H. 1988b. Nordjyllands Storke i 1988.
Riden (Dansk Orn. Foren. Nordjyllands Amt) 4:7-
10.
Skov, H. 1989a. Bolderslev storkene sat under
lup. Panurus (Dansk Orn. Foren.) 2:13-17.
Skov, H. 1989b. Storkens status i Syd- og
Sonderjylland i 1989. Panurus (Dansk Orn.
Foren.).
Skov, H. 1989c. Der Status des Weiss-Storchs in
Danemark. Pp. 55-59 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status und Schutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Skov, H. 1989d. Bolderslev storkene sat under
lup. Panurus (Dansk Orn. Foren.) 2:13-17.
Skov, H. 1989e. Bolderslev storkenes
fourageringsomrader ogfodevalg. Storkgruppen,
Dansk Orn. Foren.
Skov, H. 1989f. Bolderslev Storkenes
Fourageringsomrader og Fodelvalg. Report of the
Dansk Ornitologisk Forenings Storkgruppe,
Kobenhavn. 40pp.
Skov, H. 1990. Fremgang for de nordjyske storke.
Riden (Dansk Orn. Foren. Nordjyllands Amt) 16.
Skovgaard, P. 1937. Storken i Danmark. Danske-
Fugle 4:67-1 35.
Skyes, W.H. 1932. Catalogue of birds
(systematically arranged) of the ratorial,
gralatorial and natatorial orders, observed in
the Dukhien. Proc. Zool. Soc. Lond. 1932:149-
172.
Slater, K.R. 1958. Stray feathers. Emu 58:256.
Slater, P. 1959. Breeding periods of birds in the
Kimberley Division, Western Australia. West.
Slater, P., et al. 1970. A Field Guide to Australian
Birds, Non-Passerines. Livingstone. Wynnewood,
Pennsylvania.
Slud, P. 1964. The birds of Costa Rica,
distribution and ecology. Bull. Am. Mus. Nat.
Hist. 128.
Smirenskii, S.M. 1991a. Oriental White Stork
 action plan in the USSR. Pp. 165-178 in
Coulter, M.C., Q. Wang and C.S. Luthin
(eds), Biology and Conservation of the Oriental
White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Smirenskii, S. 1991b. Oriental White Storks
banded in the Soviet Far East. Spec. Group
Storks, Ibises, Spoonbills 4(2):4.
Smirenskii, S.M., and Ph.R. Shtilmark. 1977.
[Important habitats for the Oriental White
Stork, and Japanese and Common Crane
along the Sokhranenii River in the Priamura
Region.] Pp. 246-247 in Winter, S. (ed.) VII
All-Soviet Ornithological Conference. Zoological
Institute of the Academy of Science of the
Ukranian SSR, Kiev.
Smirenskii, S.M., and E.M. Smirenskaya. 1980.
About some rare and less studied birds of
Juish Autonomus Republic (Chabarovaki
Region). Ornithology 15:205-206.
Smirenskii, S.M., V.A. Andronov, and A.F.
Bylkov. 1987. [First results of colour marking
of the Oriental White Stork (Ciconia boyciana).]
Ornitologiya 22:194-220.
Smith, F.T.H. 1967. The Glossy Ibis near
Melbourne during 1965. Aust. Bird Watcher
3:54.
Smith, G.H. 1943. Parachute action when
settling of Sacred Ibis and Spoonbill. Ostrich
14:192-194.
Smith, K.D. 1951. On the birds of Eritrea. Ibis
93:201-233.
Smith, K.D. 1955a. The winter breeding season
of land-birds in eastern Eritrea. Ibis 97:480-
507.
Smith, K.D. 1955b. Recent records from Eritrea.
Ibis 97:65-80.
Smith, K.D. 1957. An annotated list of the birds
of Eritrea. Ibis 99:1-26, 307-337.
Smith, K.D. 1960. The passage of palaearctic
migrants through Eritrea. Ibis 102:536-544.
Smith, K.D. 1965. On the birds of Morocco. Ibis
107:493-526.
Smith, K.D. 1968. Spring migration through
southeast Morocco. Ibis 110:452-492.
Smith, K.D. 1970. The Waldrapp Geronticus
eremita (L.). Bull. Br. Orn. Club 90:18-24.
Smith, K.D., and G.B. Popov. 1953. On birds
attacking desert locust swarms in Eritrea.
Entomologist 86:3-7.
Smith, R.B., and D.R. Breininger. 1988.
Northern breeding range extension for the
Roseate Spoonbill in Florida. Fla. Field Nat.
16:65-67.
Smithers, R.H.N. 1964. A Check List of the Birds of
Bechuanaland Protectorate. Trustees Nat. Mus. S.
Rhod., Salisbury. 188pp.
Smithers, R.H.N., M.P.S. Irwin, and M.L.
Paterson. 1957. A Checklist of the Birds of
Southern Rhodesia. Rhodesian Ornith. Soc,
Salisbury.
Smuk, A. 1956-57. White Stork chatters while
flying. Aquila 63-64:329.
Smythies, B.E. 1953. The Birds of Burma, 2nd edn.
Oliver and Boyd, London.
Smythies, B.E. 1968. The Birds of Borneo, 2nd edn.
Oliver and Boyd, London.
Smythies, B.E. 1981. The Birds of Borneo, 3rd edn.
Malayan Nature Society, Kuala Lumpur.
Smythies, B.E. 1986. The Birds of Burma, 3rd edn.
S. Mattacchione and Co, Pickering, Ontario.
Snethlage, E. 1935. Beitrage zur Fortpflanzungs
biologic brasilianische Vogel.y. Ornithol. 83:1-
24, 532-562.
Snow, D.W. 1950. The birds of Sao Tome and
Principe in the Gulf of Guinea. Ibis 92:579-
595.
Snow, D.W. 1952. A contribution to the
ornithology of north-west Africa. Ibis 94:473-
498.
Snow, D.W. 1978. An Atlas ofSpeciation in African
Non-passerine Birds. Trustees of the British
Museum (Natural History), London. 390pp.
Snyder, D.E. 1966. The Birds of Guyana. Peabody
Museum, Salem, Massachusetts.
Snyder, N.F.R., and H.A. Snyder. 1969. A
comparative study of mollusc predation by
Limpkins, Everglade Kites, and Boat-tailed
Crackles. Living Bird 8:1 77-223.
Snyder, N.F.R., J.C. Ogden, J.D. Bittner, and
G.A. Grau. 1984. Larval dermestid beetles
feeding on nestling Snail Kites, Wood Storks,
and Great Blue Herons. Cowdfor86:170-174.
Sokolowski, J. 1968. Ptaki Polski. Panstwowe
Zaklady Wydawnictw Szkolnych, Warszawa.
270pp.
Soni, V.C. 1988 Ecology and Behaviour of the Indian
Black Ibis (Pseudibis paillosa Temmink).
University of Saurastra, India.
Soni, V.C., P.B. Chavda, and S. Vyas. 1989.
Feeding behaviour of the Indian Black Ibis in
various microhabitats at Rajkot and
Junagadh. Pp. 39-43 in Patel, B.H. (ed.), The
Behaviour. Proc. Nat. Symp. Anim. Behav., Sir P.P.
Institute of Science, Bhavnagar.
Sonobe, K., and N. Izawa. 1987. Endangered Bird
Species in the Korean Peninsula. The Museum of
Korean Nature and Wild Bird Society of
Japan, Tokyo.
Soothill, E., and R. Soothill. 1982. Wading Birds
of the World. Blanford Books, Poole.
Soots, R.F., Jr. and J.F. Parnell. 1975. Ecological
succession of breeding birds in relation to plant
succession on dredge islands in North Carolina. UNC
Sea Grant. Publication UNC-SG-75-27,
Willmington, North Carolina, USA.
Southern, W.E., et al. 1975. Wood Storks in
Michigan. Jack-Pine Warbler 53:158.
Souza Bazarra, J.A. d., and F. Gomez de la
Torre. 1980. Algunas notas sobre Platalea
leucorodia en Galicia. Donana Ada Vert. 7:1 1 1—
113.
Spaans, A.L. 1974. De ornithologische rijkdom
van de modderkust van Suriname. Nat.
landschap28:316-1328
Spaans, A.L. 1975a. On the present breeding
status of the Scarlet Ibis Eudocimus ruber along
the north-eastern coast of South America. Biol.
Conserv. 7:245-253.
Spaans, A.L. 1975b. The status of the Wood
Stork, Jabiru, and Maguari Stork along the
Surinam coast, South America. Ardea 63:1 16—
130.
Spaans, A.L. 1975c. Two misidentified eggs of
the American Flamingo from Surinam in the
oological collection of the Penard brothers.
Ardea 63:131-135.
Spaans, A.L. 1976a. De ligging van de reiger- en
ibiskolonies langs de kust van de Guyana's in
1976. Unpublished report, Dienst's Lands
Bosbeheer, Paramaribo.
Spaans, A.L. 1976b. Ligging reiger- en
ibiskolonies langs de kust vanSuriname in
1975. Unpublished report, Dienst's Lands
Bosbeheer, Paramaribo.
Spaans, A.L. 1977. De ligging van de reiger- en
ibiskolonies langs de kust van Suriname in
1977. Unpublished report, Dienst's Lands
Bosbeheer, Paramaribo.
Spaans, A.L. 1982a. De Rode Ibis: een
pronkjuweel van de overstromingsvlaktes van
tropisch Zuid-Amerika. Vogeljaar 30:189-193.
Spaans, A.L. 1982b. Present status of some
colonial waterbird species in Surinam, South
America./ Field Orn. 53:269-272.
Spaans, A.L. 1983a. Organochlorine
contaminants in South American Scarlet
Ibises. Unpublished report, Research Institute
for Nature Management, Arnhem.
Spaans, A.L. 1983b. Naschrift. Vogeljaar 31:157.
Spaans, A.L. 1984. Waterbird studies in coastal
Suriname: a contribution to wetland
conservation in northeastern South America.
374
Pp. 63-76 in RINM Annual Report 1983.
Research Institute for Nature Management,
Arnhem, Leersum, Texel.
Spaans, A.L. 1990a. Problems in assessing trends
in breeding populations of Scarlet Ibises and
other ciconiiform birds. Pp. 1-6 in Frederick,
P.C., L.G. Morales, A.L. Spaans, and C.S.
Luthin (eds), The Scarlet Ibis (Eudocimus
ruber) : Status, Conservation, and Recent Research,
Proc. 1st International Scarlet Ibis
Conservation Workshop, Caracas, Venezuela.
IWRB Special Publication 11, Slimbridge,
Glos, UK.
Spaans, A. (chairman). 1990b. Discussion on
developing an action plan for Scarlet Ibis
conservation in Trinidad, Guyana, Suriname,
French Guiana, and Brazil. Pp. 152-156 in
Frederick, P.C., L.G. Morales, A.L. Spaans,
and C.S. Luthin (eds), The Scarlet Ibis
(Eudocimus ruber): Status, Conservation, and
Recent Research, Proc. 1st International Scarlet
Ibis Conservation Workshop, Caracas,
Venezuela. IWRB Special Publication 11,
Slimbridge, Glos, UK.
Spaans, A.L., and B.H.J. de Jong. 1981. De
huidige stand van een aantal watervogels langs
de kust van Suriname. Panda 17:3—7.
Spaans, A.L., and B.H.J. de Jong. 1982. Present
status of some colonial waterbird species in
Suriname, South America. J. Field Orn.
53:269-272.
Spaans, A.L., M. van Wieringen, and K.
Brouwer. 1990. Scarlaken bloemen in het
donkergroene lover. De Schijnwerper gericht
op de Rode Ibis en de First International
Scarlet Ibis Conservation Workshop. Het
Vogeljaar 38:2-11.
Specialist Group on Storks, Ibises, and
Spoonbills. 1989. First Oriental Crested Ibis
raised in captivity. Spec. Group Storks, Ibises,
Spoonbills Newsletter 2(1/2): 1 1 .
Specialist Group on Storks, Ibises, and
Spoonbills. 1990. Oriental Crested Ibis. Spec.
Group Storks, Ibises, Spoonbills Newsletter
3(1/2:3.
Spector, W.S. (ed.). 1956. Handbook of Biological
Data. Saunders, Philadelphia. 584pp.
Speek, B.J. 1971. Ringverslag van het
vogeltrekstation. Limosa 44:105-132.
Speek, B.J. 1972. Ringverslag van het
vogeltrekstation. Limosa 45: 145— 1 73.
Speek, B.J. 1973a. Ringverslag van het
vogeltrekstation 191 1-1970. Limosa 46:109-
135.
Speek, B.J. 1973b. Ringverslag van het
vogeltrekstation. Limosa 46:136-165.
Speek, B.J. 1974. Ringverslag van het
vogeltrekstation. Limosa 47:63—93.
Speek, B.J. 1975. Ringverslag van het
vogeltrekstation. Limosa 48:129-157.
Spendelow, J.A., and S.R. Patton. 1988. National
atlas of coastal waterbird colonies in the contiguous
United States: 1976-1982. US Fish Wildl. Serv.
Biol. Rep. 88(5). x + 326pp, Washington DC.
Spendelow, J.A. , R.M. Erwin, and B.K.
Williams. 1989. Patterns of species co-
occurrence of nesting colonial Ciconiiformes in
Atlantic coast estuarine areas. Colonial
Waterbirds 12:51-59.
Spil, R.E., M.W. van Walstijn, and H. Albrecht.
1985. Observations on the behaviour of the
Scarlet Ibis Eudocimus ruber in Artis Zoo,
Amsterdam. Bijdr. Dierkd. 55:219-232.
Sprenger, J. 1990. Brown County's first White-
faced Ibis. Loon 62:159-160.
Sprunt, A., Jr. 1933. The Glossy Ibis in Georgia.
Auk 50:352.
Sprunt, A., Jr. 1936b. The Roseate Spoonbill in
Georgia. Auk 53:203-204.
Sprunt, A. Jr. 1936a. The Roseate Spoonbill in
South Carolina. Auk 53:75.
Sprunt, A., Jr. 1939. The present status of the
 Roseate Spoonbill in the United States. Fla.
Nat. 12(3):49-55,
Sprunt, A., Jr. 1941 a. Association of White and
Eastern Glossy Ibises. Fla. Nat. 14:74-75.
Sprunt, A., Jr. 1941b. Predation of Boat-tailed
Crackles on feeding Glossy Ibis. Auk 58:587-
588.
Sprunt, A., Jr. 1944. Northward extension of the
breeding range of the White Ibis. Auk 61:144-
145.
Sprunt, A., Jr. 1954. Florida Bird Life. Coward-
McCann, New York. 527pp.
Sprunt, A., Jr., and E.B. Chamberlain. 1949.
South Carolina Bird Life. University of South
Carolina Press, Columbia.
Sprunt, A., IV, and M.P. Kahl, Jr. 1960.
Mysterious Mycteria—Our American Stork.
Audubon Mag. 62:206-209, 234, 252.
Sprunt, A., IV, and C.E. Knoder. 1980.
Populations of wading birds and other
colonially nesting species on the Gulf and
Caribbean coasts of Mexico. Pp. 3-16 in
Schaeffer, P.P., and S.M. Ehlers (eds), The
Birds of Mexico: Their Ecology and Conservation.
Proceedings of the National Audubon
Symposium, Tiburon, California.
Sridhar, S. 1989. White and Blacknecked Storks
in the 1989 census. Newsletter Birdwatchers
29:10.
Sridharan, U. 1986. An Adjutant gobbles
ducklings. Hornbill 1986(2):26, 35.
Staav, R. 1977. Eremitibisen, en till sin existens
hotad fagelart. Fauna Flora (Stockh.) 72:39-41.
Stager, K.E. 1957. The avifauna of the Tres
Marias Islands, Mexico. Auk 74:413-432.
Stahlecker, D.W. 1989. White-faced Ibis
breeding in Rio Arriba County: second verified
nesting location for New Mexico. New Mexico
Orn. Soc. Bull. 17:2-6.
Stamper, M.N., R. Ryder, and M.M. Lett. 1972.
New nesting site for White-faced Glossy Ibis
Plegadis chihi in Colorado. J. Colo.-Wyo. Acad.
&i.7:ll.
Stangel, P.W., J.A. Rodgers, Jr., and A.L.
Bryan. 1990. Genetic variation and population
structure of the Florida Wood Stork. Auk
107:614-619.
Stangel, P.W., J.A. Rodgers, and A.L. Bryan.
1991. Low genetic differentiation between two
disjunct White Ibis colonies. Colonial Waterbirds
14:13-16.
Stark, A., and W.L. Sclater. 1906. The Birds of
South Africa. R.H. Porter, London.
Steadman, D.W., and P.S. Martin. 1984.
Extinction of birds in the Late Pleistocene of
North America. Pp. 466-477 in Martin, P.S.,
and R.G. Klein (eds), Quaternary Extinctions: a
Prehistoric Revolution. University of Arizona
Press, Tucson, Arizona.
Steele, B.B. 1980. Reproductive success of the
White-faced Ibis: effects of pesticides and
colony characteristics. M.S. Thesis, Utah
State University, Logan, Utah.
Steele, B.B. 1984. Effects of pesticides on
reproductive success of White-faced Ibis in
Utah, 1979. Colonial Waterbirds 7:80-87.
Steffen, G.L., B.K. Harris, and J. Harter. 1978.
1978 nesting season report. S. Dakota Bird Notes
30:75-77.
Stegman, B. 1962. Die verkummerte distale
Handschwinge des Vogelflugels.y. Orn.
103:50-85.
Steinbacher, G. 1937. Zur Haltung und Plege des
Schuhschnabels (Balaeniceps rex]. Gd.). Zool.
Garten (NF) 9:101-106.
Steinbacher, G. 1965. Der Storch Radolfzell BB
14412. Kosmos 61:26.
Steinbacher, G., and J. Steinbacher. 1974. Der
Bestand des WeiBstorchs Ciconia ciconia 1954-
1974 im bayerischen Regierungsbezirk
Schwaben. Ber. Naturwiss. Ver. Schwaben
1974(3/4): 75-80.
Steinbacher, J. 1936. Untersuchungen uber die
Nahrungsbiologie des Weissen Storch in
Ostpreussen 1933 und 1934. Schr. Physik. Okon.
Ges. Konigsberg 69:23-36.
Steinbacher, J. 1979. Family Threskiornithidae.
Pp. 253-268 in Mayr, E., and G.W. Cottrell
(eds), Check-list of Birds of the World, vol. 1, 2nd
edn. Mus. Comp. Zool., Cambridge,
Massachusetts.
Steinfatt, O. 1934a. Zur Brut- und
Zugverbreitung des Schwarzen und Weissen
Storches auf der Pyrenaenhalbinsel. Beitr.
Fortpfl. Vb'gel. 10:161-168.
Steinfatt, O. 1934b. Vogelbruten im Wiener
Schonbrunner Tiergarten. Beitr. Fortpfl. Vb'gel
10:57-60.
Steiof, K. 1987. Massenzug des Weiss-Storchs
(Ciconia ciconia) bei Hurghada/Aegypten.
[Very large numbers of the White Stork
(Ciconia ciconia) over Hurghada/Egypt.]
Vogelwarte 34:100-106.
Stejneger, L.H. 1884. Analecta ornithologica.
Auk 1:225-236.
Stejneger, L.H. 1887. Review of the Japanese
birds. Proc. U.S. Nat. Museum.
Stephens, F. 1904. The Roseate Spoonbill in
California. Condor 6:139.
Sterbetz, I. 1968. Assembling of molting storks in
Kardoskut. Aquila 75:29.
Sterbetz, I. 1984. K-calory demand of the heron
colonies in the Hungarian Tisza- Valley. Tiscia
(Szeged) 19:195-199.
Sterbetz, T. 1964. Glossy Ibis nesting in the
Saser Sanctuary. Aquila 69:247.
Stern, H., G. Thielcke, F. Vester, and R.
Schreiber. 1978. Rettet die Voegel-Wir Brauchen
Sie. Bund fur Umweld und Naturschutz
Deutschland, Bonn, West Germany.
Stettenheim, P. 1976. Structural adaptations of
feathers. Proc. Int. Orn. Congr. 16:385-401.
Stevenson, H.M. 1957. The relative magnitude of
the trans-gulf and circum-gulf spring
migrations. Wilson Bull. 69:39-77.
Stevenson, H.M. 1972. Records of the Scarlet
Ibis and Red-breasted Blackbird in Ecuador.
Wilson Bull. 84:99.
Stewart, R.E. 1957. Eastern Glossy Ibis nesting
in southeastern Maryland. Auk 74:509.
Steyn, P. 1963. Trio. Bokmakierie 15:5-9.
Steyn, P. 1988. Attempted kleptoparasitism of
Openbilled Storks by Greyheaded Gulls.
Ostrich 59:182.
Stinner, D.H. 1980. Estimate of wading birds in
the Macks Island Rookery, Okefenokee
Swamp, Georgia (USA), 1979. Georgia J. Sci.
138:181-183.
Stinner, D.H. 1983. Colonial wading birds and
nutrient cycling in the Okefenokee Swamp.
PhD Dissertation, University of Georgia,
Athens, Georgia.
Stoddard, D.R. 1968. The Aldabra affair. Biol.
Conserv. 1:63-69.
Stoddard, D.R. 1971. Settlement, Development
and conservation of Aldabra. Phil. Trans. Roy.
Soc.Lond.B 260:6 11-628.
Stoll, F.E. 1934. Einiges ueber den
Schwarzstorch. Ardea 23:51-56.
Stollmann, A. 1956. Die Verbreitung des WeiB-
storchs in des Umgebung der Hohen Tatra.
Vogelwarte 18:212-214.
Stollmann, A. 1958. Distribution of White Stork
at foot of High Tatra Tanap. Treat. Concerning
Tatra Natl Park 2:127-133.
Stollmann, A. 1961. Zur Verbreitung und
Okologie des Schwarzstorchs (Ciconia nigra) in
der Slowakei. Vogelwarte 21:54.
Stollmann, A. 1962. Verbreitung des
Schwarzstorchs (Ciconia nigra) in der Slowakei.
Biologia 17:365-371.
Stollmann, A. 1969. Populationsuntersuchungen
am Weiss-Storch (C. ciconia) in der Slowakei
1968. Vogelwarte 25:65-66.
375
Stollmann, A. 1975. Ciconia nigra in der Slowakei
im Jahre 1971. Ada Rerum Nat. Mus. Natl Slov.
21:231-235.
Stollmann, A. 1989. Der Weiss-Storch in der
slowakischen sozialistischen Republik. Pp. 73-
76 in Rheinwald, G., J. Ogden, and H. Schulz
(eds), Weiss-Storch: Status undSchutz, Proc. 1st
Int. Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Stone, A.C. 1913. Some swamp birds. Emu
13:82-86.
Storer, J.H. 1962. Florida's wading birds. Fla.
Nat. 35:105-106.
Storer, R.W. 1989. Notes on Paraguayan birds.
Occas. Papers Mus. Zool. Univ. Michigan, 719.
Storer, R.W. 1990. Openbill Storks. Birder's
World 4:62-63.
Storr, G.M. 1973. List of Queensland birds. W.
Austr. Mus. Spec. Publ. 5.
Stott, K., Jr. 1948. Notes on the longevity of
captive birds. Auk 65:402-405.
Stott, K., Jr. 1959. Two recent records of the
Roseate Spoonbill on the Pacific slope and
high Andes of Peru. Auk 76:244.
Stott, K., Jr. 1974. Stork Parade. Zoonooz4?:7-
11.
Stransky, D. 1929. Die Ursache des
weissfarbigen Laufes von Ciconia alba. Aquila
34-35:447.
Strazds, A. 1987. Nest of the White Stork in the
colony of Grey Heron. Putni Daba 1:97.
Street, M. and D.S. Peters. 1991. An early post
glacial record of the White Stork (C. ciconia)
from the Erfttal.J. Orn. 132:102-103.
Stresemann, E. 1927-1934. Aves. In Kukenthal,
W. (ed.), Handbuech derZoologie, Vol. 7, pp. 1—
899. Walter de Gruyter, Berlin.
Stresemann, E. 1941. Die Voegel von Celebes. J.
Orn. 89:1-102.
Stresemann, E. 1963a. Taxonomic significance of
wing molt. Proc. Int. Orn. Congr. 13:171-175.
Stresemann, E. 1963b. The nomenclature of
plumages and molts. Auk 80:1-8.
Stresemann, E. 1963c. Variations in the number
of primaries. Condor 65:449—459.
Stresemann, E., and V. Stresemann. 1966. Die
Mauser der Voegel. J. Orn. 107
(sonderheft): 1-448.
Strohl, J. 1917. Conrad Gesner's 'Waldrapp'.
Vjschr. natur. Ges. Zurich 62:501-538.
Stromar, L. 1979. A recent intensification of the
White Stork banding in Yugoslavia. Ring
9(100):62-65.
Stronach, B.W.H. 1968. The Chagana heronry in
western Tanzania. Ibis 110:344-348.
Stumpf, D. 1977. Strohhalsibis—je gehort? Bull.
Ver. Fr. Zool. Gartens Basel 38:6-10.
Stumpf, D. 1978. Zur Brutbiologie des
Strohhalsibis (Threskiornis spinicollis). Gefiederte
Welt 102:72-76.
Stunkard, H.W., and C.P. Jandal. 1965. A new
trematode, Cathaemasia senegalensis, from the
Saddle-bill Stork, Ephippiorhyncus senegalensis
(Shaw). Zoologica 50:67-73.
Sturgis, B.B. 1928. Field Book of Birds of the
Panama Canal Zone. G. P. Putnam's & Sons,
New York.
Sturm, S. 1983. Das Brutverhalten des Roten Sichlers
Eudocimus ruber im Tierpark Hellabrunn, 1982.
Geschaftsbericht 1982, Muenchener Tierpark
Hellabrunn AG, Muenchen.
Su, H., H. Liu, and S. Shen. 1989. [On the
breeding ecology of the Black Stork (Ciconia
nigra).} Acta Zool. Sin. 35:444-446.
Sun, S. 1991. Status of the storks in Lianoning
Province, China. Pp. 89-96 in Coulter, M.C.,
Q. Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Suthard, J. 1926. Wood Ibis and egret in
Hopkins Co., Kentucky. Auk 43:231-232.
Sutter, E.R. 1984. Ontogeny of the wing moult
pattern in the White Stork Ciconia ciconia. Proc.
Pan-Afr. Orn. Congr. 5:543-551.
Sutton, G.M. 1923. A visit to a Wood Ibis
colony. Bird Lore 26:391-395.
Sutton, G.M. 1967. Oklahoma Birds. University of
Oklahoma Press, Norman OK, USA.
Suzuki, S. 1980. Captive reproduction of the
Glossy Ibis. Animals Zoos 32:155-157.
Swennen, C., and E.C.L. Marteijn. 1985a.
Foraging behaviour of the Milky Stork.
Interwader Ann. Rep. 1985:27.
Swennen, C., and E. Marteijn. 1985b. Wader
feeding ecology studies in the Malay
Peninsula, Interwader Annu. Rep. 1984:13-26.
Swennen, C., and E.C.L. Marteijn. 1987. Notes
on the feeding behaviour of the Milky Stork
Mycteria cinerea. Forktail 3:63-66.
Swennen, C., and A.L. Spaans. 1985. Habitat
use of feeding migratory and local ciconiiform,
anseriform and charadriiform birds in coastal
wetlands of Surinam. Gerfaut 75:225-251.
Swinhoe, R. 1863a. On the ornithology of
Formosa. Ibis 5:416.
Swinhoe, R. 1863b. Catalogue of the birds of
China, with remarks principally on their
geographic distribution. Proc. Zool. Soc. Lond.
1863:259-339.
Swinhoe, R. 1864. Description of four new
species of Formosan birds; with further notes
on the ornithology of the island. Ibis
6(23,24):361-370.
Swinhoe, R. 1867. Jottings on birds from my
Amoy journal. Ibis 3:226-237, 385-413; 4:
52-65.
Swinhoe, R. 1871. A revised catalogue of the
birds of China and its islands, with
dcescriptions of new species, references to
former notes, and occasional remarks. Proc.
Zool. Soc. Lond. 1871:337-423.
Swinhoe, R. 1873a. On the White Stork of Japan.
Proc. Zool. Soc. Lond. 1873:512-514.
Swinhoe, R. 1873b. On the Rosy Ibis of China
and Japan. Ibis 9:249-253.
Swinhoe, R. 1873c. Untitled letter from Mr. R.
Swinhoe. Proc. Zool. Soc. Lond. 1873:729-730.
Symons, R.E. 1918. The Black Stork (Ciconia
nigra). Bull. S. Afr. Biol. Soc. 1:8-10.
Symons, R.E. 1924. The nesting of the Green
Ibis, Theristicus hagedash, in South Africa. Comp.
Ool. 1(1):10-13.
Szidat, L. 1935. Warum wirft der Storch seine
Jungen aus dem Nest? J. Orn. 83:76-87.
Szidat, L. 1940. Die Parasitenfauna des Weissen
Storches und ihre Beziehungen zu Fragen der
Okologie, Phylogenie und die Urheimat der
Storche. Z. Parasitenkd. 11:563-592,
Szidat, L. 1943. Weitere Beobachtungen uber
Parasiten und ander Krankheitserreger in aus
dem Nest geworfenen Storchen. Z. Morphol.
Oekol. Tiere 40:238-247.
Szijj, J. 1951. Gemtelpek Magyarorszagon.
[Heron colonies in Hungary.] Aquila 55—
58:81-85.
Szijj, J., and L. Szijj. 1955. Contributions to the
food-biology of the White Stork (Ciconia c.
ciconia L.). Aquila 59—62:83-94 [Hungarian
with English summary],
Taczanowski, L. 1876. Revue critique de la faune
ornithologique de la Siberie Orientale. Bull.
Soc. Zool. Fr. 1 December 1876.
Taczanowski, L. 1877. Supplement a la liste des
Oiseaux recueillis au nord du Perou occidental
par M.M. Jelski et Stolzmann. Proc. Zool. Soc.
Lond. 1877:744-754.
Taczanowski, L. 1891-93. Faune Ornithologique de
la Siberie Orientale. Memoires de I'Academie
Imperiale des Sciences de St. Petersbourg, VII Serie,
Tome 39. St. Petersburg.
Taczanowski, M.L. 1887. Liste des Oiseaux
recuellis en Coree par M. Jean Kalinowski.
Proc. Zool. Soc. Lond. 1887:596-611.
Taczanowski, M.L. 1888. Liste supplementaire
des Oiseaux recueillis en Coree par M.Jean
Kalmowski. Proc. Zool. Soc. Lond. 1888:450-
469.
Tadic, A. 1959. Cold tolerance of the White
Stork (Ciconia ciconia). Larus 11:141-147.
Takagi, N., and M. Sasaki. 1974. A phylogentic
study of bird karyotypes. Chromosoma (Berl.)
46:91-120.
Takara, T. 1979. Rare and new records of birds
from the Okinawa Islands Japan. J. Yamashina
Inst. Orn. 11:132-156.
Takashima, H. 1956a. The Japanese Stork in the
past and present. Tori 16:35-36.
Takashima, H. 1956b. Notes on the White Ibis
in Japan. Misc. Rep. Yamashina Inst. Orn. Zool.
9:16-20.
Takashima, H. 1957a. The Japanese Crested Ibis
and the White Ibis./ Fac. Set. Hokkaido Univ.
Ser. VI Zool. 13:118-124.
Takashima, H. 1957b. Notes on the Black Stork
Ciconia nigra in Japan. Misc. Rep. Yamashina's
Inst. Ornithol. Zool. 11:5-9.
Talent, G.F. 1940. Wood Ibis (Ibis ibis) in
Southern Rhodesia. Ostrich 11:49.
Tan, Y. 1986. The protected rare animals in
China. Birds: Part 2. Ciconiiformes. Chinese
Wildl. 5:5-8.
Tan, Y. 1987. The protected rare animals in
China. Birds: Part 5. Chinese Wildl. 36:29-31.
Tantzen, R. 1951. Ergebnisse der
Storchforschung im Lande Oldenburg 1949
und in den angrenzenden Gebieten. Orn. Abh.
9:11-28.
Tantzen, R. 1962. Der Weissen Storch Ciconia
ciconia (L.) im Lande Oldenburg. Oldenburger
Jahrb. 61:105-213.
Tarboton, W.R. 1968a. Check List of Birds of the
South Central Transvaal. Witwatersrand Bird
Club, Johannesburg.
Tarboton, W. 1968b. Glossy ibis on the
Witwaterstrand. Bokmakierie 20:14-15.
Tarboton, W.R. 1971. Birds of the Mosdene
Nature Reserve, Naboomspruit. S. Afr. Avif.
Ser. Percy Fitzpatrick Inst. Afr. Orn. Univ. Cape
Town 78:1-51.
Tarboton, W.R. 1977a. The status of communal
herons, ibis and cormorants on the
Witwatersrand. S. Afr.J. Wildl. Res. 7:19-25.
Tarboton, W.R. 1977b. A checklist of the birds of
the Nylsvlei Nature Reserve. S. Afr. Natn. Sci.
Propr. Rpt 15:1-14.
Tarboton, W. 1977c. The Black Stork—one of
South Africa's rarest birds. Fauna Flora
(Transv.) 29:14-15.
Tarboton, W. 1982. Breeding status of the Black
Stork in the Transvaal. Ostrich 53:151-156.
Tarboton, W., and P. Cardwell. 1968. Breeding
observations on the Black Stork (Ciconia nigra).
Bokmakierie 20(4):86-87.
Tarboton, W.R., M.I. Kemp, and A.C. Kemp.
1987. Birds of the Transvaal. Transvaal
Museum, Pretoria.
Tashian, R.E. 1953. The birds of southeastern
Guatemala. Condor 55:198-210.
Tate, A.L., and R.L. Humphries. 1980. Wood
storks nesting in Jenkins County, Georgia.
Oriole 45:34-35.
Tavakoli, E.V. 1989. The White Stork
distribution in Iran. Pp. 183-188 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status und Schutz. Proc. 1st Int.
Stork Conservation Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Taylor, D.M., C.H. Trost, and B.Jamison. 1989.
The biology of White-faced Ibis in Idaho.
West. £mfc 20:125-133.
Taylor, J.S. 1957. Notes on the birds of inland
waters in the Eastern Cape Province with
special reference to the Karoo. Ostrich 28:1-80.
Taylor, L.E. 1909. Notes from Cape Colony./ S.
Afr. Orn. Union 5:81-87.
376
Taylor, M.A., Jr. 1941. Birds from the Yucatan
Peninsula. Field Mus. Nat. Hist. Pub I. Zool. Ser.
24:195-225.
Teal, J.M. 1965. Nesting success of egrets and
herons in Georgia. Wilson Bull. 77:257-263.
Technau, G. 1965. Erfahrungen mit WeiB-
storchen bei Heuschreckenbekampfungen im
nordlichen Afrika. Vogelwarte 23:149.
Teixeira, D.M, and R.C. Best. 1981. Adendas a
ornitologia do Territorio Federal do Amapa.
Bol. Mus. Para. Emilio Goeldi 104:1-25.
Teixeira, D.M., J.B. Nacinovic, and F.B.
Pontual. 1987. Notes on some birds of
northeastern Brazil (2). Bull. Br. Orn. Club
107:151-157.
Teixeira, D.M., J.B. Nacinovic, and G. Luigi.
1988. Notes on some birds of northeastern
Brazil (3). Bull. Br. Orn. Club 108:75-79.
Teixeira, D.M., J.B. Nacinovic, and G. Luigi.
1989. Notes on some birds of northeastern
Brazil. Bull. Br. Orn. Club 109:152-157.
Teixeira, D.M., J.B. Nacinovic, and J. Dujardin.
1990. Notas sobre la distribucion y
conservacion de Eudocimus ruber (Linnaeus,
1758) en Brasil. Pp. 124-129 in Frederick,
P.C., L.G. Morales, A.L. Spaans, and C.S.
Luthin (eds), The Scarlet Ibis (Eudocimus
ruber): Status, Conservation, and Recent Research,
Proc. 1st International Scarlet Ibis Conserv.
Workshop, Caracas, Venezuela. IWRB
Special Publication 11, Slimbridge, Glos, UK.
Tejera Nunez, V.H. 1989a. Huevos de una
colonia de aves. El Rosario, Cocle, Panama.
Congr. dent. Nac. 6:114.
Tejera Nunez, V.H. 1989b. Estructura de los
nidos en aves de una colonia, El Rosario,
Cocle, Panama. Congr. dent. Nac. 6:115.
Tejera Nunez, V.H. 1990. Alimentacion en aves
de una colonia. Congr. dent. Nac. 7:151.
Tekke, MJ. 1969. Ornithologie van Nederland
1966 en 1967. Limosa 42:36-78.
Telfair, R.C., II. 1980. Additional inland nesting
records in Texas of four species of colonial
waterbirds. Bull. Tex. Orn. Soc. 13:11-13.
Telfair, R.C., II, and D.A. Swepston. 1987.
Analysis of banding and marking nestling Anhingas,
Olivaceous Cormorants, Roseate Spoonbills, ibises,
bitterns, herons and egrets in Texas. Texas Parks
and Wildlife Department Report PWD-BK-
7100-152-7/87, Austin, Texas, USA.
ten Kate, C.G.B. 1964. Ornithologie van
Nederland, 1962. Limosa 37:19-57.
ten Kate, C.G.B. 1965. Ornithologie van
Nederland, 1963. Limosa 38:24-66.
ten Kate, C.G.B. 1966. Ornithologie van
Nederland, 1964. Limosa 39:42-81.
ten Kate, C.G.B. 1967. Ornithologie van
Nederland, 1965. Limosa 40:14-58.
Terrataz, M. 1979. Turkey Project 1062. Bald
Ibis. WWFYb. 1978-1979:122.
TerresJ.K. 1980. The Audubon Society Encyclopedia
of North American Birds. A. Knopf, New York.
Thalen, D.C.P., and E.M.P. Thalen-Horst. 1972.
An observation of migrating White Storks
Ciconia ciconia (L.) in the Mesopotamian plain.
Bull. Iraq Nat. Hist. Mus. (Univ. Baghdad) 5:17-
20.
Thaler, E., E. Ettel, and S. Job. 1981. Zur
Socialstruktur des Waldrapps Geronticus
eremita—Beobachtungen an der Brutkolonie
des Alpenzoos Innsbruck./ Orn. 122:109-128.
Thauront, M. and M. Duquet. 1991.
Distribution et conditions d'hivernage de la
Cigogne Blanche, Ciconia ciconia. Av. Mali
Alanda59:\Q\-\\Q.
Thevenot, M., P. Beaubrun, R.E. Baovab, and
P. Bergieu. 1982. Compte rendu d'ornithologie
Marocaine, annee 1981. Documents de 1'Institut
Scientifique. Scientifique no. 7. Institut
Scientifique, Rabat.
Thiede, U. \982.Japanibis undJapanische Nachtigall
als Beispiele zweier Pole im Naturverstandnis der
 Japaner. Gesellschaft fuer Natur- und
Volkerkunde Ostasiens, Mitteilungen Band
90, Hamburg.
Thiemann, H. 1975. WeiBstorch-Bestand 1974 in
der Region Marrakech/Marokko. Orn. Mitt.
27:109-110.
Thienemann, J. 1910. Der Zug des Weissen
Storches (Ciconia ciconia). Zool.Jb. suppl.
12:665-686.
Thienemann, j: 1911. X. Jahresbericht (1910)
der Vogelwarte Rossitten der Deutschen
Ornithologinschen Gesellschaft. J. Orn.
59:620-707.
Thiollay, J.M. 1971. Exploitation des feux de
brausse paries oiseaux en Afrique occidentale.
Alauda 39:54-72.
Thomas, B.T. 1977. Hooding and other
techniques for holding and handling nestling
storks. N. Am. Bird Bander 2:47-49.
Thomas, B.T. 1979a. The birds of a ranch in the
Venezuelan Llanos. Pp. 213—232 in Eisenberg,
J.F. (ed.), Vertebrate Ecology of the Northern
Neotropics. Smithsonian Institution Press,
Washington, D.C.
Thomas, B.T. 1979b. Plumage succession of
nestling Maguari Storks. Bol. Soc. Venezolana
CienciasNat. 34(136):239-241.
Thomas, B.T. 1981. Jabiru nest, nest building,
and quintuplets. Condor 83:84-85.
Thomas, B.T. 1984. Maguari Stork nesting:
juvenile growth and behavior. Auk 101:812-
823.
Thomas, B.T. 1985. Coexistence and behavior
differences among the three western
hemisphere storks. Pp. 921-931 in Buckley,
P.A., M.S. Foster, E.S. Morton, R.S. Ridgley,
and F.G. Buckley (eds), Neotropical Ornithology,
Ornithological Monograph No. 36. American
Ornithologists' Union, Washington, D.C.
Thomas, B.T. 1986. The behavior and breeding
of adult Maguari Storks. Condor 88:26-34.
Thomas, B.T. 1987. Philopatry of banded
Maguari Storks and their decline in
Venezuela. Bol. Soc. Venezolana Ciencias Nat.
41:137-157.
Thomas, B.T. 1988 (1989). A comparison of the
Maguari Stork, Ciconia maguari, with the White
Stork, Ciconia ciconia. Gerfaut 78:113-119.
Thomas, B.T. 1991. A brief review of patagial
wing tags on birds. Spec. Group Storks, Ibises,
Spoonbills Newsletter 4(l):7-8.
Thomas, B.T. 1990. Nesting behavior of
sunbitterns (Eurypygahelias) in Venezuela.
Condor 92:576-581.
Thomas, W.W., Jr. 1945. Some additional
Georgia records of interest. Oriole 10:25-26.
Thompson, K. 1985. Observations on the hand
rearing of the White stork at the Bristol Zoo.
Ratel 12:71-74.
Thompson, M.C. 1966. Birds from North
Borneo. Univ. Kans. Publ. Mus. Nat. Hist.
17:377-433.
Thomson, A.L. (ed.). 1964. A New Dictionary of
Birds. Nelson, London.
Thomson, A.L. 1973. The generic name Ibis and
English term 'wood-ibis'. Ibis 115:285.
Ticehurst, C.B., P.A. Buxton, and R.E.
Cheesman. 1921-1922. The birds of
Mesopotamia./ Bombay Nat. Hist. Soc. 28:210-
250, 381-427, 650-674, 937-956.
Ticehurst, C.B., P. Cox, and R.E. Cheesman.
1926. Additional notes on the avifauna of Iraq.
J. Bombay Nat. Hist. Soc. 31:91-1 19.
Tidwell, M. 1985a. Warheads and Wood Storks.
Tidwell, M. 1985b. Wood Stork colony
threatened with loss of habitat. Bird Watcher's
Digest, Jan/Feb.
Tilson, R.L., and O.B. Kok. 1980. Habitat
ecology of Black Storks in the Kuiseb River.
Madoqua 11:347-349.
Tinbergen, L. 1934. Lepelaarsmaag. De Levende
Nat. 38:262.
Tinbergen, N. 1952. 'Derived' activities; their
causation, biological significance, origin, and
emancipation during evolution. Q. Rev. Biol.
27:1-32.
Tingay, A. and S.R. Tingay. 1976. Report on
Lake Chandala with recommendations of its
importance of a fauna reserve. W. Austr. Dept.
Fish Wildl. Rep. 26:1-30.
Tipton, I.H., and J.C. Howell. 1949. White Ibis
observed in Knox Co., Tennessee. Migrant
20:50.
Tirion,J.C. 1931. Maraboe. Trop. Natuur 20:154-
158.
Tischler, F. 1941. Die Voegel Ostpreussens, vol. 2.
Ost-Europa Verlag, Konigsberg.
Todd, W.E.C. 1948. A new booby and a new ibis
from South America. Proc. Biol. Soc. Wash.
61:49-50.
Toffoli, R. 1990. La Ciconia nera (Ciconia nigrd)
in Piemonte (Aves, Ciconiidae). Riv. Piem St.
Nat. 11:163-169.
Toffoli, R., B. Caula, and A. Pellegrino. 1988.
Caso di svernamento di Cicogna nera, Ciconia
nigra, in Piemonte. Riv. Ital. Orn. 58:213-215.
Tordoff, H.B. 1952. Note on the birds of
Jamaica. Auk 69:320-322.
Tordoff, H.B. 1956. Checklist of the birds of
Kansas. Univ. Kans. Publ. Mus. Nat. Hist.
8:307-359.
Torres Esquivias, J.A. and A.L. Claveria. 1982.
Contribucion al conocimiento de la
distribucion y biologia de la ciguena blanca
(Ciconia ciconia) en la provincia de Cordoba
(Andalucia, Esparia). Bol. Estac. Cent. Ecol.
ll(21):43-47.
Trams, E.G. 1969. Carotenoid transport in the
plasma of the Scarlet Ibis (Eudocimus ruber).
Comp. Biochem. PhysioL 28:1177-1184.
Tratz, E.P. 1960/61. Unsere gegenwartige
Kenntnis vom Waldrapp oder Klausrapp. Jb.
Ost. Arb. Kr. Wildtier-Forsch. 1.
Tree, AJ. 1978. A visit to Makgadikgadi Pan in
April 1974. Honeyguide 95:39-41.
Tree, AJ. 1982. The Black Stork in Zimbabwe.
Honeyguide 109:18-19.
Triebl, R., and H. Fruhstuck. 1979. Erhebungen
uber den Weiss-storch (Ciconia ciconia) im
Burgenland von 1963-1973. Nat. Umwelt
Burgenland, Sonderheft 2:1-31.
Tristam, H.B. 1860. On the ornithology of
northern Africa. Part 3. The Sahara. Ibis 2:68-
83.
Tristam, H.B. 1864. Report on the birds of
Palestine. Proc. Zool. Soc. Lond. 1864:426-456.
Trotignon, J. 1976. La nidification sur le Bane
d'Arguin (Mauritanie) au printemps 1974.
Alauda 44:119-133.
Trotignon, E., and J. Trotignon. 1981.
Recensement hivernal 1979-1980 des Spatules,
des Flamants et des Pelicans blancs sur le
Bane D'Arguin (Mauritanie). Alauda 49:203-
215.
Tsen-Hwang, S.A.M. 1936. The Birds of the Hopei
Province. The Vertebrates of China, vol. XV. Fan
Memorial Institute of Biology, Peking.
Tsukamoto, Y. 1991. Status of the Oriental
White Stork in Japan. Pp. 143-148 in Coulter,
M.C., Q. Wang, and C.S. Luthin (eds), Biology
and Conservation of the Oriental White Stork
(Ciconia boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Tuer, F.V. 1977. Notes on the Black Stork.
Safring News 6:21 -24.
Tuer, F.V. 1978. Notes on the Black Stork.
Honeyguide 94:15-16,18.
Tufts, W. 1949. Eastern Glossy Ibis in Nova
Scotia. Can. Field-Nat. 63:117.
Tugarinov, AJ. 1931. Die Wanderungen der
Nordasiatischen Vogel. Vogelzug 2:55-56.
377
Turner, D.A. 1978. The heronry at Lake Jipe.
Scopus 2:47-48.
Turner, M.I.M. 1964. Some observations of bird
behaviour made from aircraft in the Serengeti
National Park. Bull. Br. Orn. Club 84:65-67.
Uchida, Y. 1970. On the color change in the
Japanese Crested Ibis. A new type of cosmetic
coloration in birds. Misc. Rep. Yamashina Inst.
Orn. 6(l&2):54-72.
Uchida, Y. 1974. Feather color of Japanese
Crested Ibis. Animals Zoos 12:412-414.
Uchida, Y. 1976a. Why the Toki remained in
Sado. Anima 37:13-20.
Uchida, Y. 1976b. Taoki in the world. Anima
37:21-28.
Underwood, C.F. 1896. A list of birds collected
or observed on the lower, southern, and
southwestern slopes of the Volcano of
Miravalles and on the lower lands extending to
Bagaces in Costa Rica, with a few observations
on their habits. Ibis (ser. 7) 2:431-451.
Unitt, P. 1984. The birds of San Diego County.
Memoir 13. San Diego Society of Natural
History, San Diego, California.
Uramato, M. 1976. What does extinction mean?
Anima 37:29-36.
Urban, E.K. 1967. Possible occurrence of Whale-
headed Stork in Ethiopia./ East Afr. Nat. Hist.
Soc. Natl. Mus. 26:87-88.
Urban, E.K. 1970. Ecology of water birds from
four Rift Valley lakes in Ethiopia. Ostrich
suppl. 8:315-322.
Urban, E.K. 1974a. Flight speed and wing
flapping rate of Sacred Ibis. Auk 91:423.
Urban, E.K. 1974b. Breeding of the Sacred Ibis
Threskiornis aethiopica at Lake Shala, Ethiopia.
Ibis 116:263-277.
Urban, E.K. 1976. Studies of the avifauna of
some Rift Valley lakes of Ethiopia. Pp. 425-
436 in National Geographic Society Research
Reports, 1968 Projects.
Urban, E.K. 1978. Ethiopia's Endemic Birds.
Ethiopian Tourist Organization, Addis Ababa.
30pp.
Urban, E.K., and L.H. Brown. 1971. A Checklist
of Birds of Ethiopia. Haile Sellassie I University
Press, Addis Ababa.
Urn, AJ. 1980a. A bird sanctuary near Narora.
Newsletter Birdwatchers 20:2.
Urfi, AJ. 1980b. Birding in Aligarh. Newsletter
Birdwatchers 20:6-7.
Urfi, A J. 1988. Avifauna of Aligarh Region
(Checklist of birds and notes on habitat destruction).
Monograph, Nature Conservation Society,
Aligarh.
Urfi, AJ. 1989a. Painted Storks of Delhi Zoo.
Sanctuary 9(4):26-33.
Urfi, A J. 1989b. Breeding and habitat selection
patterns by Painted Storks Mycteria leucocephala
in Delhi Zoological Park heronries. Wader Study
Group Bull. 57:26 (abstr.).
Urfi, AJ. 1989c. Painted Stork Mycteria
leucocephala (Pennant) swallowing a snake. J.
Bombay Nat. Hist. Soc. 86:96.
Urfi, A J. 1990. Mysterious disappearance of
Painted Storks from Delhi Zoo heronries and
abrupt termination of their breeding. Newsletter
Birdwatchers 30:3-5.
Urner, C.A. 1932. Eastern Glossy Ibis in New
Jersey. Auk 49:459.
US Fish and Wildlife Service. 1983. Stork
population declines; endangered status
proposed. Endang. Species Tech. Bull. 8(3): 1, 8.
US Fish and Wildlife Service. 1984. Wood stork
listed as endangered species. Endang. Spec. Tech.
Bull. 9(3): 1,3.
US Fish and Wildlife Service. 1986. Recovery plan
for the U.S. breeding population of the Wood Stork.
U.S. Fish and Wildl. Service, Atlanta,
Georgia.
Uttley, J. 1986. Survey of coastal waterbirds in
Sulawesi Selatan March-April 1986. PHPA-
 Interwader Report No. 2. Kuala Lumpur,
Malaysia.
Uttley, J. 1987. Survey of Sulawesi Selatan to assess
the status of wetlands and to identify key sites for
breeding and migratory waterbirds. PHP A/
Interwader Report No. 2, Kuala Lumpur,
Malaysia.
Utvinenko, N.M. 1968. [On the distribution of
the Oriental White Stork in the Lower Amur.]
Ornitologiya 9:356-358.
Uys, C J. 1966. How many nest sites of the white
stork are there in South Africa? Bokmakierie
18:63.
Uys, CJ. 1968. Breeding of the White Stork
Ciconia ciconja L. at Mossel Bay, Cape. Ostrich
39:30-32.
Uys, CJ., and CJ. Broekheysen. 1966. Hadedah
(Hagedashia hagedash) nesting on telegraph pole.
Ostrich 37:239-240.
Uys, C.J., and J.G.R. Macleod. 1967. The birds
of the De Hoop Vlei region, Bredasdorp, and
the effect of the 1957 inundation over a 10-year
period (1957-1966) on the distribution of
species, bird numbers and breeding. Ostrich
38:233-254.
Uys, C J., GJ. Broekhuysen, J. Martin, and
J.G.R. MacLeod. 1963. Observations on the
breeding of the Greater Flamingo Phoenicopterus
ruber Linnaeus in the Bredasdorp District,
South Africa. Ostrich 34:129-154.
Vader, WJ.M. 1965. Bird conservation by the
'Dutch Biological Expedition Turkey 1959'.
Ardea 53:172-204.
Vaidya, A. A. 1986. Invasion of White Storks
(Ciconia ciconia) in Kachch (Kutch), Gujarat. J.
Bombay Nat. Hist. Soc. 83:433.
Valdes Miro, V. 1984. Datos de nidificacion
sobre las aves que crian en Cuba. Poeyana
282:1-27.
Valverde, J.A. 1957. Aves del Sahara Espanol.
Institute de Estudios Africanos, Madrid.
Valverde, J.A. 1958. An ecological sketch of the
Goto Dofiana. British Birds 51:1-23.
Valverde, J.A. 1959. Cuatro interesantes especies
en Andalucia. Ardeola 5:143-148.
Valverde, J.A. 1960. La nidification de la
Espatula en el sur de Espana. Ardeola 6:378.
Van der Ven, J. 1987. Asian waterfowl-1987.
Unpublished report, IWRB, Slimbridge, Glos,
U.K.
Van der Ven, J. 1988. Asian waterfowl-1988.
Unpublished report, IWRB, Slimbridge, Glos,
U.K.
Van Meter, V.B. 1985. Florida's Wood Storks.
Florida Power and Light Co., Miami.
Van Rossem, AJ. 1945. A distributional survey
of the birds of Sonora, Mexico. Occas. Pap.
Mus. Zool. La. State Univ. 21.
Van Someren, V.D. 1947. Field notes on some
Madagascar birds. Ibis 89:235-267.
Van Someren, V.G.L. 1922. Notes on the birds
of East Africa. Novit. Zool. 29:1-246.
Van Someren, V.G.L. 1956. Days with birds.
Studies of habits of some east African species.
Fieldiana Zool. 38:1-520.
Van Tets, G.F. 1978. Second amendments to the
1975 RAOU checklist. Emu 78:80-87.
Van Tyne, J., and AJ. Berger. 1959.
Fundamentals of Ornithology. John Wiley and
Sons, New York.
Vande Weghe, J.-F. 1981. L'avifaune des
papyraies au Rwanda et au Burundi. Gerfaut
71:489-536.
Vanden Berge, J.C. 1970. A comparative study
of the appendicular musculature of the order
Ciconiiformes. American Midi. Nat. 84:289-364.
Vandier, J. 1950. Une statue d'ibis. Revue
d'Egypt. 7:33-35.
Varden, J.G. 1986. Black Storks near
Featherstone. Honeyguide 32:88.
Vasvari, M. 1955. Magyarorszagi madarak
meretei. Aquila 59-62:167-184.
Vaurie, C. 1964. A survey of the birds of
Mongolia. Bull. Am. Mus. Nat. Hist. 127:103-
144.
Vaurie, C. 1965. The Birds of the Paleartic Fauna,
Non-passeriformes. H.F. and G. Witherby,
London.
Vaurie, C. 1972. Tibet and its Birds. H.F. and G.
Witherby, London.
Vergoossen, W.G. 1983. De Zwarte Ooievaar
(Ciconia nigra) in de Benelux. Veldornitol
Tijdschr. 6:39-58.
Verheijen, J.AJ. 1964. Breeding season on the
island of Flores, Indonesia. Ardea 52:194-201.
Verheugt, WJ.M. 1987. Conservation status and
action program for the Milky Stork (Mycteria
cinered). Colonial Waterbirds 10:211-220.
Verheugt, WJ.M. 1989. Status of storks in the
South Sumatra Province, Indonesia. ICBPI
IWRB Spec. Group Storks, Ibis, Spoonbills
Newsletter 2:8-9.
Verheugt, WJ.M. 1990. Conservation status and
action programme for the Milky Stork Mycteria
cinerea. Unpublished report. Kuala Lumpur.
Verheught, WJ.M., A. Purwoko, F. Danielsen,
H. Skov, and R. Kadarisman. 1991.
Integrating mangrove and swamp forest
conservation with coastal lowland
development; the Banyuasin Sembilang
Swamps Case Study, South Sumatra Province,
Indonesia. Land. Urban Plan. 20:85-94.
Verheyen, R. 1950. La Cigogne blanche dans son
quartier d'hiver. Gerfaut 40:1—17.
Verheyen, R. 1953. Exploration du Pare
National de 1'Upemba. Oiseaux Inst. Pares Nat.
Congo Beige, Fasc. 19.
Verheyen, R. 1959. Contribution a 1'anatomie et
a la systematique de base des Ciconiiformes
(Parker 1868). Bull. Inst. R. Sci. Nat. Belg.
35:1-34.
Verheyen, R. 1960. What are Ciconiiformes?
Proc. Int. Orn. Congr. 12:741-743.
Verheyen, R. 1961. A new classification for the
non-passerine birds of the world. Bull. Inst. R.
Sci. Nat. Belg. 37:1-36.
Vernon, C J. 1975. Saddlebill Stork breeding at
Rainham. Honeyguide 83:40.
Vernon, J.D.R. 1973. Observations sur quelques
oiseaux nicherus du Maroc. Alauda 41:101-
109.
Veroman, H. 1962. Vom Bestand des Weiss-
Storchs in Estland (Estnische SSR). Vogelwarte
21:291-292.
Veroman, H. 1970. Vom Bestand des Weiss-
Storchs (C. ciconia) in Estland 1962 bis 1967.
Vogelwarte 25:257-259.
Veroman, H. 1975. Ueber die Entwicklung des
Weisstorchbestandes (Ciconia ciconia) in
Estland. Ardeola 21:827-839.
Veroman, H. 1976. Distribution dynamics of the
White Stork (Ciconia ciconia) in the Baltic area.
Ornis Fenn. 53(4): 150-152.
Veroman, H. 1985. An ornithologist's
explanatory remarks. Eesti Loodus 28:515-516,
541,543.
Veroman, H. 1986. Bestandveranderungen des
Weissen Storches in Europa. Var Fagelv.
suppl. 11:225-228.
Veroman, H. 1989. Thirty-two year population
trends of the White Stork in the Estonian
S.S.R. Pp. 153-158 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status undSchutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Veroman, H. 1991. Storks in the Baltic States
and the USSR. Spec. Group Storks, Ibises,
Spoonbills Newsletter 4(1):4.
Veroman, H., and E. Schuz. 1980.
Schwarzschnabelstorch (Ciconia c. boyciana).
Vogelwarte 30:279-280.
Verschuren, J. 1978. Observations
378
ornithologiques dans les pares nationaux du
Zaire, 1968-1974. Gerfaut 68:3-24.
Verschuren, J., and A. Dupuy. 1987. Note sur les
oiseaux des Pares Nationaux littoraux du
Senegal. Gerfaut 77:405-442.
Vesey- Fitzgerald, D.F. 1955. Birds as predators
of the Red Locust. Ostrich 26:128-133.
Vesey-FitzGerald, D.F. 1959. Locust control
operations and their possible effects on the
population of the White Stork, C. ciconia.
Ostrich 30:65-68.
Vespremeanu, E. 1967. Rolul factorilor abiotici
in dinamica populatiilor de lopatar (Platalea
leucorodia) L. (Aves, Ciconiiformes) din lunca
dunarii. St. Si Cere. Biol. I. Sena Zool. 19:279-
284.
Vespremeanu, E.E. 1968a. Distribution and
biology of the spoonbill in Roumania. Ardea
56:160-177.
Vespremeanu, E. 1968b. Spoonbills in
Roumania: a correction. Ardea 56:287.
Vestjens, WJ.M. 1971. Damage to birds by
hailstorms. Emu 71:181-182.
Vestjens, W J. 1973a. Feeding of the White Ibis
on freshwater mussels. Emu 73:71—72.
Vestjens, WJ.M. 1973b. Wildlife mortality on a
road in New South Wales. Emu 73:107-1 12.
Vestjens, WJ.M. 1975a. Notes on the tracheae
and bronchi of the Australian spoonbills. Emu
75:87-38.
Vestjens, WJ.M. 1975b. Feeding behaviour of
spoonbills at Lake Cowal, NSW. Emu 75:132-
136.
Vestjens, WJ.M. 1977a. Reptilian predation on
birds and eggs at Lake Cowal, N.S.W. Emu
77:36-37.
Vestjens, WJ.M. 1977b. Status, habitats and
food of vertebrates at Lake Cowal. CSIRO
Division of Wildlife Research, Technical
Memorandum No. 12.
Vidal, A. 1977. Glossy Ibis Plegadis falcinellus on
the east Bavarian Danube. Anz. Orn. Ges.
Bayern. 16:20.
Vidal, G.W. 1879. The influence of rainfall on
the distribution of migratory waders and water
birds. Stray Feathers 8: 1 70-1 74.
Vidgen, H.G. 1921. Birds visiting Cape York
Peninsula and New Guinea. Emu 21:227-228.
Vierstein, L. 1963. Schwarzstorch (Ciconia nigra)
in Oberfranken. Orn. Mitt. 15:63.
Vigil, C. 1973. Aves Argentinasy Sudamericas.
Editorial Atlantida, Buenas Aires, Argentina.
Vincent, J. 1948. The flight of storks on
migration. Ibis 90: 151.
Vincent, J. 1966. Red Data Book, vol. 2- Aves.
IUCN, Lausanne.
Vincent, J., and G. Symons. 1948. Some notes on
the Bald Ibis, Geronticus calvus (Boddaert).
Ostrich 19:58-62.
Vincent, P.N. 1973. How to 'apple-pie' a White
Stork's nest. Bokmakierie 25:8-9.
Viney, C., and K. Phillipps. 1988. Birds of Hong
Kong, 4th edn. Government Printer, Hong
Kong. 214pp.
Vinokurov, A. A. 1965. [Ardeidae and
Threskiornithidae of the overflow lands in the
Kuban Valley and their role in fishery of the
north-western Ciscaucasia.] Rybogadn Ptitsv.
Znach.:151-155.
Visscher, M.N. d. 1976. Aspectos de manejo del
Parque Nacional Morrocoy en base a un
estudio ecologico de la avifauna. Unpublished
report, Proyecto MAC-CVG-FAO VEN 019,
Universidad de los Andes, Merida.
Visser, H.T. 1987. Een karyologische studie van
5 vogelsoorten uit de orden Ciconiiformes en
phoenicopteriformes, waarvan 4 nieuw
beschreven soorten. Doctoral scriptie,
University of Utrecht.
Vittery, A. (ed.). 1972. Bird report no. 2: 196&-
1969. Ornithol. Soc. Turkey/Sandy,
Bedfordshire, England. 208pp.
Vo Quy. 1975, Muc Luc. Chim Vietnam 1:631-
649.
Vo Quy. 1990. On the wings of peace. Natl. Hist.
Nov.:40-41.
Voeks, R., and S. English. 1981. White-faced Ibis
(Plegadis chihi) populations and distribution in the
western United States: 1979-1980. US Fish and
Wildl. Ser. Report, Portland, OR.
Vogelwarte Radolfzell. 1950. Der Storchbestand in
Sudwestdeutschland 1949. Rundbrief 1950 der
Vogelwarte Radolfzell.
Voisin, C. and J.F. Voisin. 1972. Plegadis
faldnellus. Alauda 40:39.
Volker, O. 1961. Die chemische
Charakterisierung roter Lipochrome in
GefiederderVoegel.J. Orn. 102:430-438.
Volkov, N.E. 1977. [Black Storks in the Bird
Forest Reserves.] P. 202 in Winter, S. (ed.) VII
All-Soviet Ornithological Conference. Zoological
Institute of the Academy of Science of the
Ukranian SSR, Kiev.
Vondracek,J. 1983. Zur Wiedereinburgerung
des Schwarzstorches in der CSSR. Falke
30:237-239.
Voous, K.H. 1960. Atlas of European Birds.
Thomas Nelson and Sons, New York. 284pp.
Voous, K.H. 1965. Check-list of the birds of
Aruba, Curacao, and Bonaire. Ardea 53:211-
213.
Voous, K.H. 1983. Birds of the Netherlands Antilles.
Walburg Press, Zutphen.
Voous, K.H. 1985. Additions to the avifauna of
Aruba, Curacao, and Bonaire, South
Caribbean. Pp. 247-254 in Buckley, P.A.,
M.S. Foster, E.S. Morton, R.S. Ridgely and
F.G. Buckley (eds), Neotropical Ornithology.
Ornithological Monograph 36. American
Ornithologists' Union, Washington D.C.
Vorobiev, K.A. 1954. [Birds of the Ussuri Area.}
Academy of Sciences of the U.S.S.R., Far
Eastern Branch, Moscow.
Voronov, B.A. 1985. Rare and Endangered Birds of
the Far East. Vladivostok Far East Science
Center. Valdivostok, USSR.
Vos, C. 1989. History and results of the White
Stork reintroduction program in the
Netherlands. Pp. 461-467 in Rheinwald, G., J.
Ogden, and H. Schulz (eds), Weiss-Storch:
Status undSchutz, Proc. 1st Int. Stork Conserv.
Symp., Walsrode, 1985, Schriftenr. DDA,
Braunschweig, Germany.
Vries, P.G. d. 1962. Recente ontwikkelingen in
de deltagebieden van de Rio Guadalquivir,
Zuid-Spanje. Nat. Lanschap 16:247-255.
Vries, P.G. d. 1966. De vogelkolonies van
Coppenamepunt, Suriname. Ardea 54:1-12.
Vuilleumier, F. 1985. Forest birds of Patagonia:
ecological geography, speciation, endemism,
and faunal history. Pp. 255-304 in Buckley,
P.A., M.S. Foster, E.S. Morton, R.S. Ridgely
and F.G. Buckley (eds), Neotropical Ornithology.
Ornithological Monograph 36. American
Ornithologists' Union, Washington D.C.
Wackernagel, H. 1959. Some results with colour
feeding of carotenoids in birds at the Basel
Zoological Garden. Avic. Mag. 65:20-21.
Wackernagel, H. 1964a. Brutbiologische
Beobachtungen am Waldrapp, Geronticus
eremita (L.), im Zoologischen Garten Basel.
Orn. Beob. 61:49-60.
Wackernagel, H. 1964b. Hochstalter
europaischer Vogel im Zoologischen Garten
Basel. Orn. £^.61:128-132.
Wade, P. 1982. Every Australian Bird Illustrated.
Kigby, Sydney.
Wagler,J. 1827. Systerna Avium. Cottae,
Stuttgartiae et Tubingae.
Wait, W.E. 1925. Manual of the Birds of Ceylon.
Ceylon J. Sci., Colombo.
Walker, S. 1991. Vanishing species. Illustrated
Weekly of India, Weekend Jan. 5-6:46.
Walsh, J.F. 1977. Nesting of the Jabiru Stork
(Ephippiorhynchus senegalensis) in West Africa.
Bull. Br. Orn. Club 97(4): 136.
Walsh, J.F. 1989. Wetlands of the moist-savanna
region of West Africa, and their importance to
migratory White Storks. Pp. 271-280 in
Rheinwald, G., J. Ogden, and H. Schulz (eds),
Weiss-Storch: Status undSchutz. Proc. 1st Int.
Stork Conserv. Symp., Walsrode, 1985,
Schriftenr. DDA, Braunschweig, Germany.
Walsh, J.M. 1989. Saltwater and freshwater
habitat use by Wood Storks (Mycteria
americana) at Cumberland Island National
Seashore [abstract]. Colonial Waterbird Soc.
Newsletter 13(2):26.
Walsh, J.M. 1990. Estuarine habitat use and
age-specific foraging behavior of Wood Storks
(Mycteria americana). M.S. Thesis, University of
Georgia, Athens, Georgia. 61pp.
Wang, Q. 1987. [A world-wide endangered
bird-the Eastern White Stork.] Chinese Wildl.
5.
Wang, Q. 1991. Wintering ecology of Oriental
White Storks in the lower reaches of the
Chiangjiang River, Central China. Pp. 99-106
in Coulter, M.C., Q. Wang, and C.S. Luthin
(eds), Biology and Conservation of the Oriental
White Stork (Ciconia boyciana). Savannah
River Ecology Laboratory, Aiken, South
Carolina, USA.
Ward, A.E. 1907. Birds of the provinces of
Kashmir and Jammu and adjacent districts,
part 4./ Bombay Nat. Hist. Soc. 17:943-949.
Ward, P. 1965. The breeding biology of the
Black-faced Dioch Quelea quelea in Nigeria. Ibis
107:326-349.
Warga, K. 1954. Voranzeige uber die
Erforschung der Vogelwelt des Kisbalaton.
Aquila 55-58:169-187.
Warga, K. 1955. XVIth report on bird-banding
in Hungary. Aquila 59-62:233-251.
Warncke, K. 1965. Beitrag zur Vogelwelt der
Turkey. Vogelwelt 86:1-19.
Warncke, K. 1972. Beitrag zur Vogelwelt der
Turkey im Bereich der Sudgrenze. Vogelwelt
93:23-26.
Warner, D.W., and R.M. Mengel. 1951. Notes
on the birds of the Veracruz coastal plain.
Wilson Bull. 63:288-295.
Warren, E. 1929. Ciconia alba mit weissem Lauf
in Sudafrika. Aquila 34—35:446.
Waterman, M., D. Close, and D. Condon. 1971.
Straw-necked Ibis (Threskiornis spinicollis) in
South Australia: breeding colonies and
movements. S. Aust. Orn. 26:7-11.
Waters, A. 1987. Bird usage of a heron roost in
Augusta from 1980-1987. Oriole 52:57-61.
Wayne, A.T. 1906. A contribution to the
ornithology of South Carolina, chiefly on the
coast region. Auk 23:56-68.
Wayne, A.T. 1910. Birds of South Carolina.
Contributions from the Charleston Museum 1.
Charleston Museum, Charleston, SC.
Wayne, A.T. 1922. Discovery of the breeding
grounds of the White Ibis in South Carolina.
Bull. Charleston Mus. 17:17-30.
Weber, H.H. 1938. Kafer aus einem Storchnest.
Entomol. Bl. Biol. Syst. Kaefer 34:337-338.
Weber, R. P. 1981. Viewing tower at Middle
Lake Kerang (Victoria, Australia):
preliminary results of a feasibility study. Fish
Wildl Pap. Victoria 25:1-14.
Webster, M.A. 1975. An Annotated Checklist of the
Birds of Hong Kong. Hong Kong Bird Watching
Society, Hong Kong.
Weickert, P. 1960. Reproduccion de la Espatula
en Donana en 1960. Ardeola 6:379.
Weigold, H. 1912-1913. Ein Monat Ornithologie
in den Wiisten und Kulturoasen
Nordwestmesopotamiens un Innersyriens.y.
Orn. 60:249-297, 365-410; 61:1-40.
Weidmann, U. 1971. Innate behavior. Pp. 3-12
in Dethier, V.G., R.C. Lewontin, and M.
379
Lloyd (eds), Topics in Animal Behavior, Ecology
and Evolution. Harper and Row, New York.
Wein, G.R., and W.D. McCort. 1988. Sources of
complexity in wetland mitigation. Pp. 41-50 in
Zelanzy, J. andJ.S. Feierbend, (eds),
Proceedings of Wetlands: Increasing Our Wetland
Resources. October 4-7, 1987, Washington,
D.C., National Wildlife Federation-Corporate
Conservation Council.
Weinman, A.N. 1940. Breeding of the Indian
Painted Stork under semi-domestication.
Ceylon J. Set. Sect. B Zool. 22:141-144.
Weldon, M. 1990. The management of the
European White Stork Ciconia ciconia at the
Fort Wayne Children's Zoo. AAZPA Reg. Conf.
Proc. 1990:684-687.
Weldon, W.F.R. 1883. On some points in the
anatomy of Phoenicopterus and its allies. Proc.
Zool. Soc. Lond. 1883:638-652.
Weller, M.W. 1967. Notes on some marsh birds
of Cape San Antonio, Argentina. Ibis 109:391-
411.
Weller, M.W. 1968. The breeding biology of the
parasitic Black-headed Duck. Living Bird
7:169-207.
Wells, D.R. 1972. Systematic section. Malay. Nat.
J. 25:53.
Wells, D.R. 1976. Some bird communities in
western Sabah, with distributional records,
March 1975. Sarawak Mus.J. 24:277-286.
Wells, D.R. 1985. The forest avifauna of Western
Malaysia and its conservation. ICBP Tech.
Publ. 4:213-232.
Wendnagel, A. 1937. Ein Bastard zwischen
Weiss- und Schwarzstorch. Orn. Beob. 35:1-6.
Wendnagel, A. 1949. Die Storche in der
badischen Nachbarschaft von Basel. Orn. Beob.
46:16-18.
Wendnagel, A. 1950. Die Storche in der
badischen Nachbarschaft von Basel. Orn. Beob.
47:129-130.
Wennrich, G. 1981. Paarungszeremoniell beim
Hagedasch (Hagedashia hagedash). Die Voliere
4:27-29.
Wennrich, G. 1982a. Sunbathing behaviour in
five species of ibises (Threskiornithidae) at the
Walsrode Bird Park, Germany. Avic. Mag.
88:96-100.
Wennrich, G. 1982b. Keeping Asian White
Storks Ciconia ciconia boyciana at Vogelpark
Walsrode, Germany. Avic. Mag. 88:126-129.
Wennrich, G. 1982c.Junger Waldrapp
(Geronticus eremita) beisst wahrend der
Handaufzucht sich selbst. Die Voliere 5:181-
182.
Wennrich, G. 1986. Bestandsentwicklung,
Gefahrdung und Schutz des Schwarzstorches.
Voliere 5:166-169.
Wennrich, W. 1983. Gefied Welt Grundung der
ICBP Weltarbeitgruppe fur Storche, Ibisse,
und Loffler. Seltenheiten 10:130.
Wesley, D. 1990. A few irrigation reservoirs in
Tiruchirappalli and the bird species therein
[abstract]. Pp. 53—54 in Seminar on Wetland
Ecology and Management. Bombay Nat. Hist.
Soc. Bharatpur, India.
Westernhagen, W. v. 1970. Ueber die Brutvoegel
der Bane d'Arguin (MauretanienJ.y. Orn.
111:206-226.
Westhuizen, P.M. v.d. 1967. Black Kites and
Wooly-necked Storks in South West Africa.
Ostrich 38:203.
Wetmore, A. 1926. Observations on the birds of
Argentina, Paraguay, Uruguay, and Chile.
U.S. Natl. Mus. Bull. 133:1-448.
Wetmore, A. 1931. The avifauna of the
Pleistocene in Florida (USA). Smithson. Misc.
Collect. 85:1-41.
Wetmore, A. 1943. The birds of southern
Veracruz, Mexico. Proc. U.S. Natl. Mus.
93(3164):215-340.
Wetmore, A. 1951. Four additional species for
Panama. Auk 68:525-526.
Wetmore, A. 1956. A check-list of the fossil and
prehistoric birds of North America and the
West Indies. Smithson. Misc. Collect. 131:1-105.
Wetmore, A. 1959. Birds of the Pleistocene in
North America. Smithson. Misc. Collect. 138:1-
24.
Wetmore, A. 1960. A classification for the birds
of the world. Smithson. Misc. Collect. 139:1-37.
Wetmore, A. 1962. Notes on fossil and subfossil
birds. Smithson. Misc. Collect. 145:1-17.
Wetmore, A. 1965a. Water, Prey, and Gamebirds of
North America. National Geographic Society,
Washingtpn D.G.
Wetmore, A. 1965b. The birds of the Republic of
Panama, Part 1. Smithson. Misc. Collect. 150:1-
484.
Wetmore, A., and B.H. Swales. 1931. The birds
of Haiti and the Dominican Republic. US Natl.
Mus. Bull. 155. 483pp.
Wetten, J.CJ. v. 1984. Lepelaar Platalea leucorodia
met rossige kleur. Limosa 57:28.
Wetten, J.CJ. v. 1985. A Census of the Herons,
Ibises, Spoonbills and Storks on the South and
South-east Coast of Sri Lanka in December
1984 and January 1985. Unpublished report,
Instituut voor Taxonomische Zoologie,
Amsterdam.
Wetten, J.CJ. v. 1986. The status and distribution of
the spoonbill Platalea leucorodia. Inst. Taxon.
Zool., Univ. Amsterdam. 35pp.
Wetten, J.CJ. v., and GJ.M. Wintermans. 1986.
The food ecology of the Spoonbill Platalea
leucorodia. Inst. Taxon. Zool., Univ.
Amsterdam. 59pp.
Wexler, M. 1976. Old flinthead. Natl Wildl.
14:28-32.
Wheeler, D. 1970. Observations at a small dam.
Honeyguide 64:30-31.
Wheeler, R. 1959. The R.A.O.U. Camp-out at
Noosa Heads, Queensland 1958. Emu 59:237-
249.
Whistler, H. 1918a. Notes on the birds of
Ambala district, Punjab, part 2./ Bombay Nat.
Hist.Soc. 26:172-191.
Whistler, H. 1918b. The White-necked Stork in
the Punjab./ Bombay Nat. Hist.Soc. 25:746-
747.
Whistler, H. 1937. The Vernay Scientific Survey
of the Eastern Ghats. J. Bombay Nat. Hist. Soc.
39:447-463.
Whistler, H., and N.B. Kinnear. 1949. Popular
Handbook of Indian Birds, 4th edn. Gurney and
Jackson, London. 560pp.
Whitaker,J.I.S. 1905. The Birds of Tunisia, vol. 2.
R.H. Porter, London.
White, C.M.N. 1943. Field notes on some birds
of Mwinilunga, Northern Rhodesia. Ibis
85:127-131.
White, C.M.N. 1965. A Revised Checklist of African
Non-passerine Birds. Government Printer,
Lusaka.
White, C.M.N. 1974. Three water birds of
Wailacea. Bull. Br. Orn. Club. 94:9-11.
White, C.M.N., and M.D. Bruce. 1986. The Birds
of Wailacea. British Ornithologists Union
Checklist No. 7.
White, D.H., C.A. Mitchell, and E. Cromartie.
1982. Nesting ecology of Roseate Spoonbills at
Nueces Bay, Texas. Auk 99:275-284.
Whitelaw, D. 1968. Notes on the breeding
biology of the African Spoonbill Platalea alba.
Ostrich 39:236-241.
Whitfield, A.K., and S.J.M. Blaber. 1979.
Feeding ecology of piscivorous birds at Lake
St. Lucia, pt. 2: wading birds. Ostrich 50:1-9.
Whitfield, A.K., and D.P. Cyrus. 1983. Feeding
succession and zonation of aquatic birds at
False Bay, Lake St. Lucia. Ostrich 54:65-82.
Whittall, E. 1968. The White Stork. Bokmakierie
20:88-89.
Whymper, C. 1909. Egyptian Birds. Adam and
Charles Black, London.
Wieringen, M. v., and K. Brouwer. 1990.
Morphology and ecology of the Scarlet
(Eudocimus ruber) and White Ibis (E. albus): a
comparative review. Pp. 7—15 in Frederick,
P.C., L.G. Morales, A.L. Spaans, and C.S.
Luthin (eds), The Scarlet Ibis (Eudocimus
ruber): Status, Conservation, and Recent Research,
Proc. 1st International Scarlet Ibis
Conservation Workshop, Caracas, Venezuela.
IWRB Spec. Publ. No. 11, Slimbridge, Glos,
UK.
Wieringen, M. v., K. Brouwer, and A. Spaans.
1990. Het kwetsbare leven van de rode ibis.
Panda 26:14-16.
Wiersum, K.F. 1971. De Rode Ibis. Zijn
verspreiding en bescherming in Suriname
(Oktober 1970-Januari 1971). Postgraduate
essay, Landbouwhogeschool Wageningen,
Netherlands.
Wiese, J.H. 1977. A study of the reproductive
biology of the heron, egrets, and ibis nesting of
Pea Patch Island, Delaware. Unpublished
report, Manomet Bird Observatory,
Manomet, Massachusetts.
Wigh, RJ. 1988 (1990). White Ibis on the
Piedmont Plateau. Oriole 53:26.
Wikramanayake, E.B. 1969. Some rare and
vanishing birds of Ceylon. Loris 11:374-376.
Wild Bird Society of Japan. 1982. A Field Guide to
the Birds of Japan. Wild Bird Society of Japan,
Tokyo.
Wildash, P. 1967. Third systematic list of South
Vietnam Birds, Noi son hoi. Dieu hoc Vietnam 1.
Wildash, P. 1968. Birds of South Vietnam. Chas. E.
Tuttle Co., Rutland, VT, and Tokyo.
Wilder, G.D. 1934. Migration notes. China J.
21:135-139.
Wilder, G.D. 1940. Migration notes. China J.
32:199-204.
Wilder, G.D., and H.W. Hubbard. 1924. List of
the birds of Chihli Province./ N. China Branch
R. Asiatic Soc. 55:156-239.
Wilder, G.D., and H.W. Hubbard. 1926. List of
the birds of Chihli Province with additions and
corrections./ N. China Branch R. Asiatic Soc.
57:184-194.
Wilder, G.D., and H.W. Hubbard. 1938. Birds of
Northeastern China. Peking Natural History
Bull., Handbook no. 6, Peking, China. 700pp.
Wilkins, W.J., and C. Wilkins. 1990. An
Abdim's Stork with a broken leg. Mirafra 7:
8-9.
Williams, H.C. 1913. White Ibis (Guara alba) in
Missouri. Auk 30:268.
Williams, J.G. 1964. A Field Guide to the Birds of
East and Central Africa. Hough ton Mifflin,
Boston.
Williams, J.G. 1967. A Field Guide to the National
Parks of East Africa. Collins, London.
Williams, J.G., and N. Arlott. 1980. A Field Guide
to the Birds of East Africa. Collins, London.
Williams, M.D. 1986a. Report on the Cambridge
Ornithological Expedition to China 1985.
Unpublished report, Cambridge University.
Williams, M.D. 1986b. Preliminary report on the
China Cranewatch 1986. Unpublished report,
Cambridge University.
Williams, M.D. 1971. White Ibis in Giles
County. Migrant 42:58.
Williams, M.D., D.N. Bakewell, G.J. Carey, and
SJ. Holloway. 1986. On the bird migration at
Beidaihe, Hebei Province, China, during
spring 1985. Forktail 2:3-20.
Williams, M.D., S J. Holloway, S.R.B.
Thompson, W. Hsu, and L.Jin. 1991. The
migration of Oriental White Storks Ciconia
bqyciana and Black Storks Ciconia nigra at
Beidaihe Beach, China. In Coulter, M.C., Q.
Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
380
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Williamson, WJ.F. 1916. The Giant Ibis
(Thaumatibisgigantea).J. Nat. Hist. Soc. Siam
2:71-72.
Williamson, WJ.F. 1921. The Giant Ibis
(Thaumatibisgigantea) in Cambodia./ Nat.
Hist. Soc. Siam 4:96.
Wilson, A., and C.L. Bonaparte. 1831. American
Ornithology. Edinburgh.
Wilson, P.J. 1957. Breeding of African spoonbill
Platalea alba (Scop.) in the South Western
Cape. Ostrich 28:236.
Wilson, R.T.A. 1983. Saddlebill Stork. Redwing
30-31.
Wilson, R.T. 1988. Birds of the Sahel Zone in
Central Mali. ICBP Tech. Publ. 7:171-180.
Wilson, S.A., and G. Allport. 1985. Milky Stork
(Ibis cinereus) and birds of the Java plain. Bull.
Oriental Bird Club 1:12-14.
Winter, S. (ed.). 1977. VIIall-Soviet Ornithological
Conference. Zoological Institute, Academy of
Science of the Ukranian SSR, Kiev.
Winter, S.W. 1978b. The breeding biology of the
Oriental White Stork, Ciconia boyciana Swinhoe,
in the Middle Amur Region. Tr. Zool. Inst.
Akad. Nauk SSSR 76:9-23.
Winter, S.W. 1982. Nesting of the Black-billed
White Stork (Ciconia boyciana Swinhoe) in the
middle Amur Region. Pp. 75-100 in Gavrilov,
V.M., and R.L. Potapov (eds), Ornithological
Studies in the USSR, vol. 1. USSR Academy of
Sciences, Moscow.
Winter, S.W. 1991. Diet of the Oriental White
Stork (Ciconia bqyciana Swinhoe) in the middle
Amur region, USSR. In Coulter, M.C., Q.
Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Winterbottom,J.M. 1936. Distributional and
other notes on some Northern Rhodesian
birds. Ibis(ser. 75)6:763-791.
Winterbottom, J.M. 1937. The birds of the dams
in the eastern province of Northern Rhodesia.
Ostrich 8:68-83.
Winterbottom, J.M. 1939. Miscellaneous notes
on some birds of Northern Rhodesia. Ibis (ser.
14) 3:712-734.
Winterbottom, J.M. 1958. African Spoonbill
nesting colony on the Karoo. Ostrich 29:89.
Winterbottom, J.M. 1960. Report on the Cape
Bird Club Vlei Counts, 1952-58. Ostrich
31:135-168.
Winterbottom, J.M. 1964a. An expedition to the
Etosha Pan Game Reserve. Bokmakierie 16:
18-20.
Winterbottom, J.M. 1964b. Notes on a journey
to and from Etosha Pan, August—September
1963. Ostrich 35:327-329.
Winterbottom, J.M. 1965a. Notes from
Bushmanland. Ostrich 36:133.
Winterbottom, J.M. 1965b. Avifaunal
relationships between the Neotropical and
Ethiopian regions. Hornero 10:209-214.
Winterbottom, J.M. 1966. The birds of the
Okavango Valley. Results of the Percy
Fitzpatrick Institute of African Ornithology-
Windhoek States Museum Joint Expeditions 3.
Winterbottom, J.M. 1968. A checklist of the land
and fresh water birds of the Western Cape
Province. Ann. S. Afr. Mus. 53:1-276.
Winterbottom, J.M. 1969. Water birds in
Ovamboland. Ostrich 40:27-28.
Winterbottom, J.M. 1971. A Preliminary Checklist
of the Birds of South West Africa. SWA Scientific
Society, Windhoek.
Winterbottom, J.M. 1972. Range of the
Hadedah. Ostrich 43:186.
Winterbottom, J.M. 1977. The White Stork in
South Africa, 1974-75. Ostrich 48(3&4): 116-
118.
Winterbottom, J.M. 1979. A Guide to the Birds of
the S. W. Cape. Cape Bird Club, Cape Town.
Witherby, H.F., F.C.R. Jourdain, N.F.
Ticehurst, and B.W. Tucker. 1939. The
Handbook of British Birds, vol. 3. H.F. and G.
Witherby, London. 399pp.
Wittmann, J. 1932. Das Storchsterben am
Kamberg in den Drakensbergen Sudafrikas.
Vogelzug 3:140-141.
Wittmann, U., and H. Ruppert. 1984.
Kunstliche Erbrutung und Aufzucht des
Waldrapp, Geronticus eremita (L., 1758). Zoo/.
Garten (NF) 54:427-438.
Wodzicki, K. 1934. Studies on the Stork in Poland.
II. The Stork in Polish Silesia. Krakow.
Wojciechowski, Z., and E. Ogrodowczyk. 1978.
Rozmieszczenie i efektywnosc legow bociana
bialego (Ciconia ciconia) w wojewodztwie
miejskim lodzkim w porownaniu z materialami
z obszaru bylych powiatow Lowicz i Leczyca.
Notatki Orn. 19(l-4):27-38.
Wolfer, S. 1979. Der Schwarzstorch im sudlichen
Burgenland. Nat. Umwelt Burgenland2:14~-76.
Wolff, R.L. 1953. Wood Ibis in Tyrell County,
N.C. Chat 17:92.
Wolff, WJ. 1966. Spring and summer
observations from Mesolonghion, Greece.
Ardea 54:68-75.
Wolff, WJ. 1967. Watervogeltellingen in het
gehele Nederlandse Deltagebied. Limosa
40:216-225.
Wollert, H. 1969. Recent trends in the existence
of storks in the teterow area. Falke Maonatsschr
Orn. Vivarienkunde Ausg. A. 16:120-12.
Wolley, J. 1846. The spoonbill in Andalucia.
Zoologist 4:1213-1214.
Wolstenholme, P. 1961. Hadedahs accept
humans. Afr. Wildlife 15(3):245-250.
Won, H.G. 1966. Black-faced Spoonbill and its
protection. Korean Nat. 1966:8-11.
Won, P.O. 1963. The urgent problem of bird
protection in Korea with special suggestions
for habitat improvement and wildlife
management. ICBP Bull. 9:121-129.
Won, P.O. 1967. The present status of some
threatened and rare birds in Korea, 1962-
1966. ICBP Bull. 10:109-110.
Won, P.O. 1971. The status of bird conservation
in Korea, 1966-1970. ICBP Bull. 11:242-247.
Won, P. 1973. Wildlife and Flowering Plants. Korea
Background, Korea Overseas Information
Service, Seoul.
Wong, P.L., and R.C. Anderson. 1986. Revision
of the genus Desportesius Chabaud and
Campagna 1949 [Nematoda: Acuarioidea)
mainly from the gizzard of ciconiiform birds.
Can.J. Zoo/. 64:2520-2530.
Woo, H.C., S.W. Kim, and P.-O. Won. 1961.
Avifauna of Korea. Pp. 1-19 in Avi-mammalian
Fauna of Korea. Inst. Agr., Suwon.
Wood, C.A. 1923. The Cayenne or River Ibis in
British Guiana. Condor 25:199-201.
Wood, D.S. 1983. Phenetic relationships within
the Ciconiidae (Aves). Ann. Carnegie Mus.
52:79-112.
Wood, D.S. 1984. Concordance between
classifications of the Ciconiidae based on
behavioral and morphological data. J. Orn.
125:25-37.
Wood, D.S., and R.C. Leberman. 1987. Results
of the Carnegie Museum of Natural History
Expeditions to Belize. III. Distributional notes
on the birds of Belize. Ann. Carnegie Mus.
56:137-160.
Woods, R.W. 1975. The Birds of the Falkland
Islands. Anthony Nelson, Shrops, UK.
Woolfenden, G.E. 1959. A Pleistocene avifauna
from Rock Spring, Florida. Wilson Bull.
71:183-187.
World Birdwatch. 1988. Ibises in the 'Obo'.
World Birdwatch 10(3-4):5.
World Birdwatch. 1989. Dwarf Ibis: big news.
World Birdwatch 11(4):5.
Wray, D.L., and H.T. Davis. 1959. Birds of North
Carolina. N. C. Dept. Agr., Raleigh. 434pp.
Wrege, P.H. 1980. Social foraging strategies of
White Ibis, Eudocimus albus. PhD Dissertation,
Cornell University, Ithaca, New York.
Wright, A.H., and F.H. Harper. 1913. A
biological reconnaissance of Okefenokee
Swamp: the birds. Auk 30:477-505.
Wright, W.B., and R. Fowler. 1955. (Plegadis
falcinellus) breeding record for New Jersey.
Audubon Field Notes 9:370.
Wu, Z., X. Han, X. Li, and L. Wang. 1991a. A
large migratory population of the Oriental
White Stork in China. Pp. 67-74 in Coulter,
M.C., Q. Wang, and C.S. Luthin (eds), Biology
and Conservation of the Oriental White Stork
(Ciconia boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Wu, Z., L. Wang, S. Wu, and Q. Yang. 1991b.
Handrearing and releasing Oriental White
Storks. Pp. 181-192 in Coulter, M.C., Q.
Wang, and C.S. Luthin (eds), Biology and
Conservation of the Oriental White Stork (Ciconia
boyciana). Savannah River Ecology
Laboratory, Aiken, South Carolina, USA.
Wu, Z., X. Han, L. Wang. 1991c. Reobservation
on the autumn grouping of Oriental White
Storks. Chinese Wildl. 59:12-14.
Wunderlich, L. 1886. Beitrage zur
vergleichenden Anatomic und
Entwickelungsgeschichte des unteren
Kehlkopfes der Vogel. Nova Acta Ksl. Leop.-
Carol. Dtsch. Akad. Naturforscher4S(l):\^80.
W.W. Brehm-Fonds. 1983. The most endangered
ibis in the world. Flying Free 1(2).
W.W. Brehm-Fonds. 1984. A major Crested Ibis
conservation effort in China. Flying Free
2(2):5-7.
W.W. Brehm-Fonds. 1985. Help for the
endangered Oriental Crested Ibis. Flying Free
3(1-2):11.
Wylie, S.R. 1982. Breeding the African Spoonbill
Platalea alba at the St. Louis Zoological Park,
U.S.A. Avic. Mag. 88:70-71.
Wyndham, E. 1978. Birds of the Milparinka
district and Cooper Creek Basin. Emu 78:179-
187.
Yaacob, N., S.H. Yatim, and D. Parish. 1989.
Milky Storks in Malaysia. Spec. Group Storks,
Ibis, Spoonbills Newsletter 2:9.
Yamashina, Y. 1961. Birds in Japan. Tokyo News
Service, Tokyo. 233pp.
Yamashina, Y. 1962a. History and present status
of Japanese Crested Ibis Nipponia nippon
(Temminck). ICBP Bull. 8:77-78.
Yamashina, Y. 1962b. Present status of the
Japanese White Stork. ICBP Bull. 8:99-102.
Yamashina, Y. 1963. Endangered species in
Japan. ICBP Bull. 9:88-90.
Yamashina, Y. 1967. The status of endangered
species in Japan. ICBP Bull. 10:100-109.
Yamashina, Y. 1967. The plight of the Japanese
Crested Ibis. Animals 10:275-277.
Yamashina, Y. 1969. Japanese crested ibis —
protection and propagation programme on
Sado Island, Japan. WWF Yb. 1968:147-148,
240.
Yamashina, Y. 1977a. The status of endangered
species in Japan. ICBP Bull. 8:100-103.
Yamashina, Y. 1977b. [The present status of the
Japanese Crested Ibis Nipponia nippon and the
Japanese White Stork Ciconia ciconia boyciana in
continental Asia.] Misc. Rep. Yamashina Inst.
Orn. 9:131-142.
Yamashina, Y. 1978. The feeding of Japanese
Crested Ibises. Pp. 161-164 in Temple, S.A.
(ed.), Endangered Birds: Management Techniques
for Preserving Threatened Species. University of
Wisconsin Press, Madison.
Yamashina, Y., and S. Takano. 1959. Report on
381
the Japanese White Stork census. Misc. Rep.
Yamashina Inst. Orn. 6:505-521.
Yamasita, C. and M. dePaula V. 1986. Sobre
Anilhamento e migragao de Mycteria americana
no Pantanal. // Enav, Rio de Janeiro.
Yan, F. 1991. A preliminary study on the
wintering ecology of White Storks. Pp. 129-
134 in Coulter, M.C., Q. Wang, and C.S.
Luthin (eds), Biology and Conservation of the
Oriental White Stork (Ciconia boyciana).
Savannah River Ecology Laboratory, Aiken,
South Carolina, USA.
Yang, L. 1984. The Crested Ibis. PRC Quarterly,
April.
Yepez Tamayo, G. 1955. El order Ciconiiformes
y sus representantes en Venezuela. Mem. Soc.
Cienc. Nat. La Salle 15:5-44.
Yepez Tamayo, G. 1963. Ornitologia de las islas
Margarita, Coche y Cubagua (Venezuela).
Segunda parte. Mem. Soc. Cienc. Nat. La Salle
23:167-249.
Yonetani, Y. 1981. News from Ueno Zoo, Tokyo.
Japanese Crested Ibis update. Anim. Keeper's
For. 8:204.
Yong, L. 1984. The Crested Ibis. PCC Quarterly,
April.
Yoshi, M. 1971. The present status of the
Japanese Crested Ibis and the Japanese White
Stork. ICBP Bull. 11:168-169.
Young, C.G. 1928. A contribution to the
ornithology of the coastland of British Guiana.
Ibis(ser. 72)4:748-781.
Young, E., and S.P. Kruger. 1966. Fatal
infestation of the Bald Ibis Geronticus calvus
with Tropisurus americanus. J. S. Afr. Vet. Med.
Assoc. 37:240.
Young, V.E., and Pence, D.B. 1979. (Neottialges
(Pelecanectes) ibisicola) sp. N. (Acari:
Hypoderidae) from the subcutaneous tissue of
the White-faced Ibis (Plegadis chihi)
(Ciconiiformes: Threskiornithidae). J.
Parasitol. 65:659-661.
Zablocka, T. 1959. Spring arrival of the White
Stork, Ciconia ciconia (Linn.), in Poland in
1946-1952. Acta Orn. 5:283-299.
Zahl, P.A. 1950. Search for the Scarlet Ibis in
Venezuela. Natl. Geogr. Mag. 97:633-661.
Zahl, P. A. 1954. Coro-Coro, The World of the Scarlet
Ibis. Bobbs-Merrill Co., New York.
Zahl, P.A. 1967. New scarlet bird in Florida
skies —and a pink one too. Natl. Geogr. Mag.
132:874-882.
Zapata, A.R.P., and H.S. Martinez. 1972.
Algunas aves citadas y otras poco frequentes
para el sur de la Provincia de Buenos Aires.
Acta Zoo/. Lilloana 29:181-199.
Zar, J.H. 1968. Standard metabolism
comparisons between orders of birds. Condor
70:278.
Zarza, A.C., and J. Araujo. 1972. Neuvas
observaciones de Platalea leucorodia en el
interior. Ardeola 16:253-254.
Zhang, R. 1991. World's first Crested Ibis bred
from artificial incubation. Bird World 12:20-21.
Zhang, Z.-Y. 1986. Gem of the Orient. Living
Bird ft. 5:28-30.
Zhao, S. et al. 1990. Cathaemasiahians of Black
Stork (Ciconia nigra). Chinese Wildl.
1990(2): 125.
Zimmer, J.T. 1938. Notes on migration of South
American birds. Auk 55:405-410.
Zimmermann, D. 1955. Ein schwimmender
WeiBstorch. Orn. Beob. 52:60-62.
Zink, G. 1963a. Populationsuntersuchungen am
Weissen Storch in SW-Deutschland. Proc. Int.
Orn. Congr. 13:812-818.
Zink, G. 1963b. Der WeiBstorch-Bestand in
Baden-Wurttemberg, 1960-1962. Beitr.
Naturkd. Forsch. Suedwestdtschl. Beih. 22:89-96.
Zink, G. 1967. Populationsdynamik des Weissen
Storchs in Mitteleuropa. Proc. Int. Orn. Congr.
14:191-215.
Zlotorzycka,J. 1959. [Mallophaga of wild and
captive storks.] Ada Parasitol. Pol. 7:171-177.
Zollick, H. 1986. Ergebnisse der vierten
internationalen Bestandserfassung vom WeiB-
storch im Bezirk Rostock 1984. Falke 33:82-83.
Zollick, H. 1987. Juvenile mortality of White
Storks (Ciconia ciconia) in Rostock, Germany.
Beitr. Vogelkde. 33:276-280.
Zook-Rimon, Z., and A. Dofan. 1989a.
Preliminary observations on the Glossy Ibis
(Plegadis falcinellus) on the Akko Plain. Isr. J.
Zool. 36:59.
Zook-Rimon, Z., and Dotan. 1989b. The effect of
fire on a breeding population of the Glossy Ibis
(Plegadis falcinellus) at Mishmar Hayam. Isr.J.
Zool. 36:154-155.
Zotta, A. 1940. Lista sobre el contenido
estomacal de las aves Argentinas. Hornero
7:402-411.
Zotta, A.R., and S. da Fonseca. 1937. Sinopsis de
382
los Ciconiiformes Argentinos. Hornero 6:395—
418.
Zuppke, U. 1974. Ueberwinternder WeiBstorch
in der Elbaue. Falke 21:248.
Zusi, R.L. 1984. A functional and evolutionary
analysis of rhynchokinesis in birds. Smithson.
Contrib. Zool. 395.
Zylva, T.S.U. d. 1969. The Spoonbill. Loris
11:377-378.
 Index
Page numbers in bold type refer to the main entries for each species
Abdim's Stork (Ciconia abdimii) 9, 11, 29, 70, 71, 73, 74,
75-9, 82, 84, 85, 88, 102, 137, 291
Adjutant Stork see Greater Adjutant Stork; Lesser Adjutant
Stork
African Openbill Stork (Anastomus lamelligerus) 11, 29, 31,
60,61,62,63-7, 166,289
African Spoonbill (Platalea alba) 24, 25, 28, 166, 214, 255,
256, 267-70, 277, 280, 326
American White Ibis (Eudocimus ruber) 13, 17, 20, 21, 22,
31, 32, 34, 37, 146-56, 161, 163, 164, 166, 170, 251,
306,318
American Wood Stork (Mycteria americana) 17, 21, 30, 31,
32, 33, 34, 35-42, 47, 50, 51, 93, 121, 284
Anastomus lamelligerus 11, 29, 31, 60, 61, 62, 63-7, 166, 289
A. oscitans 11, 18, 23, 25, 26, 27, 45, 55, 59-62, 65, 66, 67,
105,219,288
Asian Openbill Stork (Anastomus oscitans) 11, 18, 23, 25,
26, 27, 45, 55, 59-62, 65, 66, 67, 105, 219, 288
Australian White Ibis (Threskiornis molucca) 6, 13, 215,
217, 218, 219, 221-5, 227, 228, 229, 300
Balaeniceps rex 7, 9, 10, 14, 30, 32, 135, 139-45, 305
Bald Ibis (Geronticus calvus) 17, 20, 29, 34, 73, 236-40,
321
Barefaced Ibis (Phimosus infuscatus) 24, 94, 157-61, 164,
170,307
Black Ibis (Pseudibispapillosa) 16, 19, 20, 164, 241-4, 245,
247, 320
Black Stork (Ciconia nigra) 11,13, 16, 17, 20, 68-74, 75,
77, 79, 82, 83, 84, 94, 95, 111, 290
Blackfaced Spoonbill (Platalea minor) 14, 16, 257, 261,
262-6, 325
Blacknecked Stork (Ephippiorhychus asiaticus) 12, 17, 19, 21,
24, 31, 33, 106, 109-13, 115, 118, 122, 123, 166, 224,
299
Bostrychia carunculata 20, 196-8, 314
B. hagedash 20, 22, 29, 32, 65, 191, 192-5, 197, 201, 202,
203,205,237,243,313
B. h. brevirostris 193, 194, 195
B. h. erlangeri 195
B. h. hagedash 193, 194
B. h. nilotica 193, 194
B. olivacea 16, 191, 197, 199-202, 203, 205, 206, 315
B. o. akleyorum 199,201
B.o.bocagei 199,201,202
B. o. cupreipennis 199, 201, 202
B. o. olivacea 199,201
B. o. rothschildi 199, 210, 202
B. rara 20, 32, 191, 197, 201, 203-6, 316
Buffnecked Ibis (Theristicus caudatus) 20, 32, 173, 177, 179,
181,183-7,311
Cercibis oxycerca 159, 176-8, 310
Ciconia abdimii 9, 11, 29, 70, 71, 73, 74, 75-9, 82, 84, 85,
88, 102, 137,291
C. boyciana 11, 16, 24, 74, 94, 95, 102, 103-8, 298
C. ciconia 11, 17, 19, 21, 24, 33, 34, 50, 60, 69, 70, 71, 72,
73, 74, 77, 78, 79, 94, 96-102, 103, 105, 106, 107-8,
137, 296-7
C. c. asiatica 97, 98, 100, 102
C. c. boyciana 102
C. c. ciconia 97, 98
C. episcopus 11, 12, 34, 70, 72, 74, 75, 77, 78, 80-5, 87, 88,
89, 90, 95, 102, 292-3
C. e. episcopus 90
C. e. neglecta 90
C. maguari 2, 11, 34, 37, 74, 79, 85, 89, 91-5, 102, 121,
122, 123, 161,295
C. nigra 11,13, 16, 17, 20, 68-74, 75, 77, 79, 82, 83, 84,
94,95, 111,290
C. stormi 1, 11, 16, 27, 74, 83, 84, 85, 86-90, 102, 294
Crested Ibis see Madagascar Crested Ibis; Oriental Crested
Ibis
Ephippiorhynchus asiaticus 12, 17, 19, 21, 24, 31, 33, 106,
109-13, 115, 118, 122, 123, 166, 224, 299
E: a. asiaticus 114
E. senegalensis 12, 18, 21, 24, 31, 33, 106, 112, 113,
114-18, 122,300
Eudocimus ruber 13, 17, 20, 21, 22, 31, 32, 34, 37, 146-56,
161, 163, 164, 166, 170, 251, 306, 318
E. r. albus 149
E. r. ruber 149
Eurasian Spoonbill (Platalea leucorodia) 3, 17, 20, 26, 213,
252-7, 259, 261, 263, 264, 265, 267, 269, 322-3
Geronticus calvus 17, 20, 29, 34, 73, 236-40, 321
G. eremita 1, 16, 17, 19, 20, 29, 230-5, 319
Giant Ibis (Thaumatibisgigantea) 14, 16, 164, 245-7, 321
Glossy Ibis (Plegadisfalcinellus) 2, 13, 65, 159, 162-7, 170,
171, 172, 177, 189, 227, 231, 241, 308
Greater Adjutant Stork (Leptoptilos dubius) 16, 19, 24, 123,
126, 127,128-32, 135, 138,303
Green Ibis (Mesembrinibis cayennensis) 32, 159, 164,
188-91,312
Hadada Ibis (Bostrychia hagedash) 20, 22, 29, 32, 65, 191,
192-5, 197, 201, 202, 203, 205, 237, 243, 313
384 Index

Harpiprion caerulescens 179-82, 310 P. leucorodia 3, 17, 20, 26, 213, 252-7, 259, 261, 263, 264,
265, 267, 269, 322-3
Jabiru mycteria 12, 18, 21, 24, 32, 37, 93, 112, 119-23, 301 P. L archeri 253, 255, 257
Jabiru Stork (Jabiru mycteria) 12, 18, 21, 24, 32, 37, 93, P. L balsaci 253, 255, 257
112,119-23,301 P. L leucorodia 255, 257
P. L regia 257
Leptoptilos crumeniferus 8, 22, 34, 78, 112, 116, 125, 126, P. minor 14, 16, 257, 261, 262-6, 325
127, 129, 130, 131, 132, 133-8, 141, 144, 304 P. regia 7, 14, 25, 166, 223, 256, 257, 258-61, 264, 265,
L. c. cristata 209 266,267,271,273,280,324
L. c. urschi 207, 209 Plegadis chihi 13, 159, 161, 164, 166, 168-72, 173, 189, 309
L. dubius 16, 19, 24, 123, 126, 127, 128-32, 135, 138, 303 P.falcinellus 2, 13, 65, 159, 162-7, 170, 171, 172, 177, 189,
L.javanicus 16, 45, 46, 57, 123, 124-7, 129, 130, 131, 135, 227,231,241,308
302 P. f. peregrinus 16 7
Lesser Adjutant Stork (Leptoptilos javanicus) 16, 45, 46, 57, P. ridgwayi 13, 172, 173-5, 309
123, 124-7, 129, 130, 131, 135, 302 Plumbeous Ibis (Harpiprion caerulescens) 179-82, 310
Lophotibis cristata 16, 202, 207-10, 316 Pseudibispapillosa 16, 19, 20, 164, 241-4, 245, 247, 320
P.p.davisoni 241,243,244
Madagascar Crested Ibis (Lophotibis cristata) 16, 202, P. p. papillosa 241,243
207-10,316 Puna Ibis (Plegadis ridgwayi) 13, 172, 173-5, 309
Maguari Stork (Ciconia maguari) 2, 11, 34, 37, 74, 79, 85,
89, 91-5, 102, 121, 122, 123, 161, 295 Roseate Spoonbill (Platalea ajaja) 14, 29, 274, 275-81,
Marabou Stork (Leptoptilos crumeniferus) 8, 22, 34, 78, 112, 328
116, 125, 126, 127, 129, 130, 131, 132, 133-8, 141, Royal Spoonbill (Platalea regia) 7, 14, 25, 166, 223, 256,
144, 304 257, 258-61, 264, 265, 266, 267, 271, 273, 280, 324
Mesembrinibis cayennensis 32, 159, 164, 188-91, 312
Milky Stork (Mycteria cinerea) 16, 43-7, 51, 55, 105, 219, Sacred Ibis (Threskiornis aethiopicus) 5, 6, 17, 21, 34, 65, 78,
285 194, 211-15, 217, 219, 221, 223, 224, 229, 239, 269,
Mycteria americana 17, 21, 30, 31, 32, 33, 34, 35-42, 47, 50, 317
51,93, 121,284 Saddlebill Stork (Ephippiorhynchus senegalensis) 12,18,21,
M. cinerea 16, 43-7, 51, 55, 105, 219, 285 24, 31, 33, 106, 112, 113, 114-18, 122, 300
M. ibis 11, 22, 25, 46, 48-52, 53, 65, 98, 116, 117, 214, Sharptailed Ibis (Cercibis oxycerca) 159, 176-8, 310
286 Shoebill (Balaeniceps rex) 7, 9, 10, 14, 30, 32, 135, 139-45,
M. leucocephala 22, 30, 43, 45, 50, 51, 53-7, 60, 61, 62, 305
127,287 Spotbreasted Ibis (Bostrychia rara) 20, 32, 191, 197, 201,
203-6,316
Nipponia nippon 16, 19, 21, 59, 218, 248-51, 321 Storm's Stork (Ciconia stormi) 1,11,16, 27, 74, 83, 84, 85,
86-90, 102, 294
Olive Ibis (Bostrychia olivacea) 16, 191, 197, 199-202, 203, Strawnecked Ibis (Threskiornis spinicollis) 4, 13, 166, 218,
205,206,315 223,224,226-9,319
Openbill Stork see African Openbill Stork; Asian Openbill
Stork Thaumatibis gigantea 14, 16, 164, 245-7, 321
Oriental Crested Ibis (Nipponia nippon) 16, 19, 21, 59, 218, Theristicus caudatus 20, 32, 173, 177, 179, 181, 183-7, 311
248-51,321 T. c. branickii 183, 185, 186
Oriental White Ibis (Threskiornis melanocephalus) 6, 28, 30, T. c. caudatus 183, 185, 186
46, 166,213,216-20,223,318 T. c. hyperorius 183, 185, 186
Oriental White Stork (Ciconia boyciana) 11, 16, 24, 74, 94, T. c. melanopsis 183, 185, 186
95, 102, 103-8, 298 Threskiornis aethiopicus 5, 6, 17, 21, 34, 65, 78, 194, 211-15,
217, 219, 221, 223, 224, 229, 239, 269, 317
Painted Stork (Mycteria leucocephala) 22, 30, 43, 45, 50, 51, T.a.abbotti 211,213,214,215
53-7,60,61,62, 127,287 T. a. aethiopicus 5,211,215
Phimosus infuscatus 24, 94, 157-61, 164, 170, 307 T.a.bernieri 5,211,213,214,215
P. i. berlepschi 157, 158 T. melanocephalus 6, 28, 30, 46, 166, 213, 216-20, 223, 318
P. i. infuscatus 157, 158, 159 T. molucca 6, 13, 215, 217, 218, 219, 221-5, 227, 228, 229,
P. i. nudifrons 157, 158 300
Platalea ajaja 14, 29, 274, 275-81, 328 T. m. molucca 225
P. alba 24, 25, 28, 166, 214, 255, 256, 267-70, 277, 280, T. m. pygmaeus 221, 223, 225
326 T. m. strictipennis 225
P.flavipes 8, 14, 43, 166, 260, 261, 271-4, 280, 327 T. spinicollis 4, 13, 166, 218, 223, 224, 226-9, 319
 Index 385

T. s. faroi 229
Waldrapp Ibis (Geronticus eremita) 1, 16, 17, 19, 20, 29,
230-5, 319
Wattled Ibis (Bostrychia carcunculata) 20, 196-8, 314
White Ibis see American White Ibis; Australian White Ibis;
Oriental White Ibis
White Stork (Ciconia ciconia) 11,17, 19, 21, 24, 33, 34, 50,
60, 69, 70, 71, 72, 73, 74, 77, 78, 79, 94, 96-102, 103,
105, 106, 107-8, 137, 296-7
see also Oriental White Stork
Whitefaced Ibis (Plegadis chihi) 13, 159, 161, 164, 166,
168-72, 173, 189,309
Wood Stork, American 17, 21, 30, 31, 32, 33, 34, 35-42,
47, 50, 51, 93, 121, 284
Woollynecked Stork (Ciconia episcopus) 11,12, 34, 70, 72,
74, 75, 77, 78, 80-5, 87, 88, 89, 90, 95, 102, 292-3
Yellowbilled Spoonbill (Plataleaflavipes) 8, 14, 43, 166,
260, 261, 271-4, 280, 327
Yellowbilled Stork (Mycteria ibis) 11, 22, 25, 46, 48-52,
53, 65, 98, 116, 117, 214, 286