TA F BIOGEO

Fzlndamen~als of ~ ~ o g e o g ~presents
a ~ ~ y an engaging and comprehensive introduction to biogeography,
explaining the ecology, geography, and history of animals and plants. Defining and explaining the nature
of populations, communities and ecosystems, the book examines where different animals and plants live
and how they came to be living there; investigates how populations grow, interact, and survive, and how
communities are formed and change; and predicts the shape of communities in the twenty-first century.

Illustrated throughout with informative diagrams and attractive photos (many in colour), and including
guides to further reading, chapter summaries, and an extensive glossary of key terms, Fzlndamentals
of ~ ~ o g e o g ~ a ~ ~ explains
clearly y key concepts, life systems, and interactions. The book also tackles the most
topical and controversial environmental and ethical concerns including: animal rights, species exploitation,
habitat fragmentation, biodiversity, metapopulations, patchy landscapes, and chaos.

F,mdamentals o f ~ ~ o g e o g presents
~ a ~ ~ y an appealing introduction for students and all those interested in gaining
a deeper understanding of the key topics and debates within the fields of biogeography, ecology and the
environment. Revealing how life has and is adapting to its biological and physical surroundings,
Huggett stresses the role of ecological, geographical, historical and human factors in fashioning animal and
plant distributions and raises important questions concerning how humans have altered Nature, and how
biogeography can affect conservation practice.

Richard John Huggett is a Senior Lecturer in Geography at the University of Manchester

 ROUTLEDGEFUNDAMENTALS OF PHYSICAL
GEOGRAPHY SERIES
Series Editor: John Gerrard

Thls new series of focused, introductory textbooks presents comprehensive, up-to-date introductlons to the
fundamental concepts, natural processes and humadenvironmental impacts wlthin each of the core physical
geography sub-disciplines: Biogeography, Climatology, Hydrology, Geomorphology and Soils.

Uniformlydesigned, each volumeincludesstudent-friendly features: plentifulillustratlons, boxed case

studies, key concepts and summaries, further reading guides and a glossary.
 FUNDAMENTALS OF
BIOGEOGRAPHY

Richard John Huggett

Routledge Fundamentals of Physical

Geography

London and New York

 First published 1998
by Routledge
11 New Fetter Lane. London EC4P 4EE

Simultaneously published In the USA and Canada

by Routledge
29 West 35th Street, New York, NY 10001

0 1998 Richard John Huggett

The right of Richard John Huggett to be identified as the Author of thls Work has been asserted by
him In accordance wlth the Copyrlght, Deslgns and Patents Act 1988

Typeset In Garamond by Keystroke, Jacaranda Lodge, Wolverhampton

Prinred and bound in Great Brltaln by The Bath Press

All rights reserved. N o part of thls book may be reprlnted or reproduced

or utilized in any form or by any electronic,
mechanical, or other means, now known or hereafter
Invented, including photocopying and recording, or In any
Information storage or retrieval system, without permmion
in wrltlng from the publishers.

Brrtrrh Library Cutulo~rringm P~bliratronData

A catalogue record for this book IS available from the Brltish Library

Library of Conpess Cutalopng 111 Prrbiirutron Data

Huggett, Rlchard J.
Fundamentals of hiogeographyiRlchard John Huggetr
p. cm. - (Routledge Fundamentals of Physlcal Geography Serles)
Includes bibliographical references and index
1. Biogeography. I. Title. 11. Series
Q H 8 4 . H 8 4 1998
5 7 8 ' . 0 9 4 c 2 1 97-38998

ISBN 0-415-15498-7 (hbk)

ISBN 0-41 5-1 5499-5 (pbk)
For m y family
This Page Intentionally Left Blank
 vii

CONTENTS

Serzes editor's pre/ke ix

Lzst o f plates xi
...
Lzst of jgures x111
List of tables xvi
List o f Boxes xvii
Author's pr&z xix

INTRODUCTION: STUDYING BIOGEOGRAPHY 1

1 W H A T IS BIOGEOGRAPHY? 4
2 LIFE AND THE ENVIRONMENT 13
3 THE DISTRIBUTION OF ORGANISMS 43
4 POPULATIONS 84
5 INTERACTINGPOPULATIONS 109

6 COMMUNlTIES 138

7 COMMUNITY CHANGE 171

8 LIFE, HUMANS, AND MORALITY 209

226
235
240
254
This Page Intentionally Left Blank
 S E R I E S EDITOR’S PREFACE

W e are presently livlng In a time of unparalleled change and when concern for the envlronment has never
been greater.Globalwarmingandclimatechange, possible rlsing sea levels, deforestatlon, desertlfication
and wldespread soil erosion are just some of the issues of current concern. Although 1 t is the role of human
activity in such issues that is of most concern, this actlvlty affects the operation of the natural processes that
occur wlthin the physical environment. Most of these processes and their effects are taught and researched
within the academic disclpline of physlcal geography. A knowledge and understandingof physlcal geography,
and all it entails, is virally Important.
It IS the aim of this Fvndamentuls of Physzrul Geogrupby Serm to provide, In five volumes, the fundamental
nature of the physical processes that act on or just above the surface of the earth. The volumes In the series
are Climatology, Geomorphology,Biogeography, Hydrology and Soi1.r. The topics are treated in sufficient breadth
and depth to provide the coverage expected in a Fundunmtuls series. Each volume leads Into the toplc by out-
lining the approach adopted. Thls is important because there may be several ways of approaching lndivldual
topics. Although each volume is complete initself, there are many explicit and implicit references to the
toplcs covered in the other volumes. Thus, the five volumes together provide a comprehenslve inslght into
the totality that is Physlcal Geography.
The flexibility provided by separate volumes has been deslgned to meet the demand created by the variety
of courses currently operating In hlgher educatlon instltutlons. The advent of modular courses has meant that
physical geography I S now rarely taught, in its entirety, in an ‘all-embraclng’ course but I S generally split into
its main components. This is also the case with many Advanced Level syllabuses. Thus students and teachers
are being frustrated lncreaslngly by lack of suitable books and are having to recommend texts of which only
a small part mlght be relevant to their needs. Such texts also tend to lack the detail required. I t is the aim
of this series to provide lndivldual volumes of sufficient breadth and depth to fulfil new demands. The
volumes should also be of use to sixth form teachers where modular syllabuses are also becomlng common.
Each volume has been written by hlgher educatlon teachers wlth a wealth of experlence In all aspects of
the toplcs they cover and a proven ability in presentlng information in a lively and lnterestlng way. Each
volume provldes a comprehensive coverage of the sub~ect matter using clear text divided into easily accessible
sectlons and subsectlons. Tables, figures and photographs are used where appropriate as well as boxed case
X SERIES EDITOR'S PREFACE

studies and summary notes. References to important prevlous studies and results are included but are used
sparingly to avoid overloading the text. Suggestlons for further reading are also provided. The m a n target
readership is introductory level undergraduatestudents of physical geography or envlronmental science,
but there will be much of Interest to students from other disclplines and I t is also hoped that slxch form
teachers will be able to use the lnformatlon that I S provided In each volume.
John Gerrard, 1997
 PLATES

COLOUR

All colour plates appear in two plate sections between pp. 60-1 and pp. 156-7

1 Yellow-footed rock wallaby (Petrogale xanthrop/r.r)

2 Puma o r mountain lion (Felis concolor)
3 A cycad (Zatttia iitdenii) In Ecuador
4 Common cassowary (Casuurirrscas/urru.r)
5 Common wallaroo or euro (Macropus robustus)
6 Leadbeater’s possum (Gyrttnobelideus feadbeateri)
7 Honey possum or noolbender (Tarszpes rostratm)
8 Cattle egrets ( B / h I c u s ibis)
9 (a) Canadian beaver (Cartor tanadetrsr.r)(b) Beaver dam and lodge, Rocky Mountains Natlonal
Park, Colorado
10 (a) Red squlrrel (Scrurns zwlgarrs) (b) Grey squmel (Scrwrds rarofinenszs)
1 1 Sklpper butterfly (He.rperra conmu), on the North Downs, Surrey
12 (a) Retlculated velvet gecko (Oedrrra retzcxfrta) (b) Glmlet gum (Eucalyptrrs .ralrrbris) in a
wheat crop, near Kellerberrin, Western Austkalia
13 Golden lion tamarin (Leontidens rosalia)

BLACK A N D WHITE

1.1 Central Americm or Baird’s taplr (Tdprr1r.r bairdi), Belize 8

1.2 Nestor parrots (a) Kaka (Ne.rtor v/Nrrzdiorrulz.r)(b) Kea (Nestor notabi1i.r) 9
2.1 Splnifex hopplng mouse (Notoray.r rrlexis) - world champion urlne concentrator 26
3.1Tuatara (Sphenotlvnpzrnctatrrs) 50
xii PLATES

3.2 Brown or spotted kiwi (Aptwix awtralis) 72

3.3 A relict Antillean insectivore (Solenodon) 74
3.4 Muntjac deer (Muntzacus reevesi) 80
5.1 Ratel or honey badger (Mellivma capensis) 111
7.1 Plant succession at Glacier Bay, Alaska (a) Upper Muir inlet (b) Goose Cove (c) Muir Pomt
(d) York Creek 176
7.2 Secondary succession in Great Basm ghost towns, United States (a) Terrill (b) The sole
remaining building (c) Wonder (d) The foundanon at Wonder 180
7.3 Successional pathways on Rakata (a) Casualina equiserifolta woodland (b) Lowland Neonauclea
calycina forest ( c ) Lowland Ficus pubinwvis-Neonauclea calycma forest (d) Upland forest 183
7.4 Semi-permanent prairie wetland, Stutsrnan County, North Dakota 2 00
 FIGURES

1.1 Breeding distribution of the ring ouzel 7

1.2 Tapirs: their origin, spread and present distribution 9
1.3 (a) Sclater and Wallace classification of faunal regions (b) Numerical classification of
mammal distributions 11
2.1 Tolerance range and l i m m 16
2.2 Ecological valency, showmg the amplitude and position of the optimum 17
2.3 Temperature control in poikilotherms and homeotherms 19
2.4 Temperatures at an Esklmo dog's extremities 20
2.5 Lethal ambient temperatures for four populations of woodrats, western United States 21
2.6 Examples and explanation of climate diagram 24
2.7 Ecozones of the world 29
2.8 Soil and vegetation toposequences on Hodnet Heath, Shropshire 34
2.9 Plant life-forms 38
2.10 Proportion of plant life-forms in various ecozones 40
2.1 1 Autoecological accounts for the bluebell in the Sheffield region 41
3.1 Romanian hamster: a species with an endemlc and restricted distribution 44
3.2 Two restricted and endemic plant families: Degeneriaceae and Leitneriaceae 46
3.3 Two widespread plants: sunflower and grass families 46
3.4 Zonal climatlc distributions of four plant families 47
3.5 Protea, banksia, and grevillea family, and magnolia and tulip tree family 48
3.6 Alpine marmot: a climatic relict 49
3.7 (a) Scattergraph of the greatest north-south versus the greatest east-west dimension of North
American snake specles ranges (b) Some shapes and sizes of ranges 52
3.8 Home ranges (a) Land Iguana, Sante Fe (b) Red fox, Unlversity of Wisconsin arboretum 54
3.9 Boreal forest in Canada (a) Predicted forest types (b) Observed forest types 57
3.10 Predicted and observed distributions of longleaf pine and Florida poison tree 58
3.1 1 Winter distributlon and abundance of the eastern phoebe 59
xiv FIGURES

3.12 How spermatophyte flora reached Rakata, Krakatau Islands, 1883-1989 60

3.13 Widest ocean gaps crossed by terrestrlal animals 61
3.14 Bird abundance on various islands between New Gulnea and New Brltaln 62
3.15 Westward spread of the European starling in North Amerlca 64
3.16 History of South American mammals 65
3.17 Yapok or water opossum 66
3.18 Convergent evolutlon of the marsuplal sabre-tooth and placental sabre-tooth 67
3.19 Glyptodont: a tanklike Pleistocene herblvore descended from old xenarthran Invaders 68
3.20 Unlque South Amerlcan mammals that evolved from Late Cretaceous condylarthran Invaders 68
3.21 Bolivar Trough and Panamanlan Isthmus 69
3.22 Distribution of Me.ra.ra//r//.r in the Permlan period 70
3.23 Dlstributlon of Lystrusu~/r~~.r In the Early Triassic perlod 71
3.24 Geological history of the Caribbean region over the last 100 million years 75
3.25 Dlstributlon of Amerlcan mlnk and munt~ac deerIn Britain 79
3.26 Effect of the Indian mongoose on Pac~ficisland lizards 81
3.27 Effect of the chestnut blight in Watershed 41, western North Carolina 82
4.1 Mam areas occupled by the red deer populatlon, Lyme Park, Cheshlre 85
4.2 Exponential growth of a hypothetlcal population 87
4.3 Logistlc growth of a hypothetical population 87
4.4 Internally driven populatlon dynamlcs: stable points, stable cycles and chaos 90
4.5 Population crash in tlcklegrass: hlgh seed and low seed denslty monocultures 91
4.6 Survivorshlp curves of Dall mountaln sheep, Amerlcan robin and Brltish robin 93
4.7 Granville fritillary population in &and, Finland 96
4.8 Ecologlcal strategles in plants: competitor, stress tolerator and ruderal 102
4.9 The great auk's former distribution 105
4.10 Shrlnklng range of the Amerlcan blson 106
5.1 Two unpalatable Trinidadian butterflies resemble one another: Mullermn mimics 112
5.2 Distributions of native red squirrel and introduced grey squirrel in Britain 115
5.3 Vertical distributlon of larvae and adult acorn barnacles on intertldal-zone rocks, Millport,
Scotland 117
5.4 Warblers in spruce forests, Maine, United States 118
5.5 Beak size of ground finches on the Galipagos Islands 119
5.6 Herbivore-plant interactions 126
5.7 Ant-eatlng mammals 127
5.8 Cycles populatlon
Canadian
snowshoe
inand
density
129
lynx
of
hare
5.9 Observed cycles In Ala~okl,
prey,
western
predators
Finland
130
and
5.10 Outcomes of Gause's laboratory experments wlth protozoans as prey and
predator
131
5.11 Population cyclesin a laboratoryexperiment:apredatorymite feeds uponanothermite132
5.12 Populationchanges inalaboratoryexperlment:azukibean weevil andtwoparasitic wasps 133
6.1
Hertfordshlre
Wood,
Great
Northaw ( a ) Geology (b) Soils 139
6.2
distributlon
Tree Hertfordshire
Wood, InGreat
Northaw 1.40
6.3 Productlon zones, consumption zones and blogeochemlcal cycles In ecosystems 143
biomes different
6.4 Distribution
in of blomass 144
6.5 Mean net prlmary
production for the biospherecomponent
its
and biomes
145
6.6
patternWorldprlmary net of terrestrlal production 146
 FIGURES xv

6.7 Grazing and detrital feeding relations in an ecosystem, showmg commlnutlon spirals 148
6.8 (a) Carbon cycle (b) Nitrogen cycle (c) Phosphorus cycle (d) Sulphur cycle 150
6.9 Carbon stored in blomass, litter, humus and charcoal in the major ecozones 152
6.10 A food web, Wytham Wood, Oxfordshire 153
6.1 1 Ecological pyramids: primary producers and a plant-parasite-hyperparasite food chain 153
6.12 Sea otter distribution 155
6.13 What holds communitles together? (a) 'The world is green' hypothesls (b) 'The world is
prickly and tastes bad' hypothesis ( c ) 'The world I S whlte, yellow and green' hypothesis 157
6.14 Simple and complex food webs: repercusslons of removlng trophic levels 158
6.15 DDT biomagnlfication in the Long Island estuary food web, New York 160
6.16 Specles-area curve for Hertfordshire plants 161
6.17 (a) Herpetofauna (amphibians plus reptiles) diversity on West Indian islands (b) Bird species
on Scottish ~slands 162
6.18 Spec~es diverslty,island area and habltat diverslty, Shetland 164
6.19 Latitudinal diversity gradient mammal species richness in the Amerlcds 165
6.20 Controls on biotic diversity In polar, temperate and tropical zones 167
6.2 1 Latitudinal variations in abiotic pressure and biotic pressure actlng on spec~esof three
hypothetical body plans 168
7.1 Types of climax communities 172
7.2 Positlons of glacier t e r m m and Fastie's study sltes, Glacier Bay, Alaska 175
7.3 Location of the Krakatau island group 177
7.4 Secondary succession in Cedar Creek Natural History Area, Minnesota 179
7.5 Indivldualistic response of small North American mammals smce Late Quaternary 184
7.6 Land-cover transformatlon, A D 900-1977 187
7.7 Skipper butterfly decline in England during the twentieth century 188
7.8 Location of the Santee River and proposed river diversion 190
7.9 Bottomland forest community subjected to annual flood durations 192
7.10 Habitat changes in the Santee River study area 193
7.1 1 Atchafalaya Delta and Terrebonne Parish marshes study area, southern Louisiana 194
7.12 Observed distribution of habitats in the Atchafalaya-Terrebonne study area 195
7.13 Location of North American pralrie wetlands 199
7.14 Changes in biome area predicted by the MAPPS model under various scenarios 201
7.15 Geographical shift In blomes induced by global warming 202
7.16 Ecotone changes induced by global warming 203
7.17 Simulations of forest biomass dynamics over one millennium in response to climatic change
induced by increasing levels of carbon dioxide in eastern North Amerlca 204
7.18 Simulated changes in species composltion of forests In eastern North America 207
8.1 Integration of ecologlcal, economlc and social needs in a decision-analysls model 220
 TABLES

1.1 Blogeographical regions and subregions, as defined by Alfred Russel Wallace 12

2.1 Habitat scales 14
2.2 Temperature tolerance in plants 22
3.1 Species classed according to range size and cosmopolitanism 45
3.2 Mean geographlcal ranges of Central and North American mammal species grouped by order 51
4.1 Life table of a hypothetical population 92
4.2 Parameters for a cohort-survival model of the blue whale (Balaenoptwa musculus) population 94
4.3 Demographic parameters for a wandering albatross (Diomedza exdans) population 95
4.4 Environmental contingencies and ecological strategies in plants 100
4.5 A key for identifylng ecological strategies of herbaceous plants 101
5.1 Interactions between population pairs 110
5.2 Herbivorous tetrapods 122-3
5.3 Main pest control technlques 134
6.1 Mammals In Northaw Great Wood, Hertfordshlre 141
6.2 Human appropriatlon of net prlmary production in the 1980s 147
6.3 The diversity of livmg thmgs 161
6.4 Habitat classification used in Shetland study 164
6.5 Some factors thought to Influence species diverslty gradients 166
7.1 Area occupied by each habitat type for three years for which data are available 196
8.1 Brands of environmentalism and their characteristics 214
8.2 Slogans for deep and shallow ecology 216
8.3 Different attitudes of deep and shallow ecologrsts 2 17
 BOXES

1.1 What's in a name? Classifylng organlsms 6

2.1 The electromagnetic spectrum emltted by the Sun 18
2.2Reptiles In deserts 25
2.3Mammals in deserts 26
2.4' Plant life-forms 38-9
3.1Accounting for regularities inrange size 53
3.2AcurlousSouthAmericanmarsupial 66
4.1 The red deer population In Lyme Park, Cheshlre, England 85-6
5.1 The hawthorn and the American robln: a frult-frugwore system 124
5.2 Populatlon models of herblvore-plantsystems I 26-7
6.1 Removingtrophic levels - three ideas 157
6.2 Latitudinal diversity gradients, In genera, fiamilies and orders 168-9
7.1 Key features of FORFLO 191
7.2Amphibiansand global warmlng 198
8.1Deep ecology 2 16-17
8.2 Religlon, Nature and the World Wildlife Fund 218
This Page Intentionally Left Blank
 AUTHOR'S PREFACE

Biogeographymeansdifferentthingstodifferentpeople. To biologists, it is traditionallythehlstoryand

geography of animals (zoogeography) and plants (phytogeography). Thls historical biogeography explores the
long-term evolution of life and the Influence ofcontlnental drift, global climatic change, and other large-scale
environmental factors. Its origins lie In seventeenth-century attempts to explain how the world was restocked
by animalsdisembarklngfromNoah'sark.Itsmodernfoundatlonswere l a l d by CharlesDarwlnand
Alfred Russel Wallace in the second half of the nineteenth century. The sclence of ecology, which studies
communities and ecosystems, emerged as an dependent study in the late nineteenth century. An ecolog~cal
element then crept into traditlonal biogeography. It led to analytical and ecologlcal biogeography. Analytlcal
biogeography considers where organisms live today and how they disperse. Ecological biogeography looks at
the relations between life and the environmental complex. It used to consider mainly present-day conditlons,
but has edged backwards into the Holocene and Plelstocene.
Physlcal geographers have a keeninterestinbiogeography.Indeed,somearespeclalistteachersinthat
field. Biogeography courses have been popular for many decades. They have no common focus, their content
varyingenormouslyaccordingtotheparticularinterests of the teacher.However,many courses show a
preference for analytical and ecological biogeography, and many include human impacts as a major element.
Biogeography is alsobecominganImportantelementinthegrowingnumberofdegreeprogrammes
in environmental sclence.Biogeography courses In geographyandenvironmental science departments are
supported by agood range offine textbooks.Popularworksinclude B~ogeograph~': N'ztwal nnd C h d
(Simmons 1079),B L ~ . Bio<qeogrlrphy
~IC (Pears 1085), Btogeogr@5): A S t d y ofP1~~wt.rI N rhr Ecarphrrr (Tivy 1992),
~ ~N:E ~ a l o g ~ cm~ r/ ld .E'twl//tton~lryApproach (Cox and Moore l093), the last being in Its fifth
and B i o p ~ q p p h A
editlon with a slxth In preparation.
As therc I S no dearth of excellent textbooks, why is I t necessary to write a new one! There are at least
four good rtxsons for dolng so. First, all the popcllar texts, though they have been reissued as new editlons,
have a 1970s a l r about them. It IS a long time slncc a hasrc biogeography text appeared that took a fresh,
up-to-date.andgeographlcally focused look at thesuhlect.Second,humaninteractlonwlthplantsand
anlmals 1s now a central theme In geography, In environmental sclence, and in environmentalb~ology.
Exlstlng textbooks tackle thls topic, but therc is much more to he said about applicatlon of biogeographlcal
xx AUTHOR’S PREFACE

and ecologlcalideas In ecosystem management. Thlrd, novel Ideas In ecology are gulding research In bio-
geography. It I S difficult to read articleson ecologicalbiogeographywlthoutmeetingmetapopulations,
heterogeneous landscapes, and complexity. None of these toplcs is tackled in existlng textbooks. They are
difficulttoplcstostudyfrom research publications because theycontainformldabletheoreticalaspects.
Nevertheless, it is very important that students should be familiar with the baslc Ideas behind them. First-
year and second-year undergraduates can handle them if they are presented in an Informative and lnterestlng
way that avoids excessive mathematlcalformalism.Fourth,environmentalism in itsglorlousvariety has
mushroomed Into a vast Interdisciplinary ~uggernaut. It impinges on btogeography to such an extent that I t
would be inexcusably remiss not to let it feature in a substantlal way. It is a facet of biogeography that geog-
raphy students find fascinating. Without doubt, a blogeography textbook for the next millennium should
~ncludediscussion of envlronmental and ethical concerns about such pressing ISSUCS as species exploltatlon,

mvironmental degradation, and blodiversity. However, biogeographyis a vast sub~ectand all textbook wrlters
adopt a somewhat lndivldualistlc vlewpolnt. This book I S no exception. It stresses the role of ecological,
geographical, hlstorlcal, and human factors in fashlonlng animal and plant distributlons.
I s h o ~ ~ llike
d to thank many people who have made the completion of thls book possible. Nick Scarle
patiently drew all the diagrams. Sarah Lloyd at Routledge bravely took yet another Huggett book on board.
Several peq’le kindly provlded me with photographs. Rob Whltraker and Chrls Fastleread and Improved the
section on vegetatlon succession. Michael Bradford and other colleagues In the Geography Department at
ManchesterIJniversity didnotinterruptmysabbatlcalsemester too frequently.DerekDavenportagam
discussed a l l manner of Ideas with me. And, as always, my wife and family lent their willing support.

Rlchard Huggett
Poynton
December 1997

Note
Every effort has been made to trace the owners of all copyright material. In a few cases, the copyright owners
could not be traced. Apologies are offered to any copyright holders whose rights may have been unwittingly
infrlnged. The copyright of photographs remalns with the Individuals who supplied them.
 INTRODUCTION:
STUDYINGBIOGEOGRAPHY
Biogeography deals wlth the geography,ecology, and
history of life - where it lives, how it lives there,
and how i t came to live there.It has threemain
branches - analytical blogeography, ecological bio-
geography,and historical biogeography.Historical
biogeography considers the influence of contlnental
drift,globalclimaticchange,andother large-scale
environmental factors on the long-term evolution of
life. Ecological biogeography looks at the relations
between life andtheenvironmental complex. Ana-
lytical biogeography examines where organisms live
today and how they spread. It may be considered as
a division of ecological biogeography.
This book explores the ecological and historical
btogeography of anlmals and plants, and,in doing so,
i t considers human lnvolvement in the livlng world.
It is designed to lead students through the malnareas
of modern biogeographlcal investigation. Baslc Ideas
arecarefullyexplained usmgnumerous examples
from around the world. The chief polnts are summa-
rized at the endof each chapter. Each chapter also has
essay questlons, whlchare designedtoconsolidate
key ideas, andsuggestions for further reading. The
glossarywill helpstudentstounderstand technical
terms.
Chapter 1 addresses the baslc question:What is
biogeography,Chapter 2 looks at how life copes
with Its biological and physical environments.It
covers four areas. First, it exammesthe places in
which organisms live, focusing on habitats (landscape
elements, landscapes, and reglons), habitatrequire-
ments(habltatgeneralists,habltat specialists), and
ecological tolerance (limiting factors, tolerance range,
ecological valency). Second, it discusses climatlc
factors - radiation and light, temperature, molsture
(including snow), and climatic zones (ecozones, bio-
mes, zonoblomes, and oroblomes). Third, it explalns
how substrate and soil, topography (altitude, aspect,
Inclination,insularity),anddisturbance influence
livingthmgs.Fourth, it looks at ways of livlng
(ecological niches, ecological equlvalents), life-forms,
and autoecological accounts.
Chapter 3 Investigates the distributlon of organ-
isms - where they live and how they came to live
there.It deals with five topics. First, it illustrates
the geographlcal patterns displayed by species (and
higher-orderunits, such as families) - large and
small, widespread andrestricted(micro-endemics,
endemics, pandemlcs, and cosmopolites), contmuous
and broken (evolutlonary, jump dispersal, geological,
andclimaticdisjunctions), relict groups, the geog-
raphy of range sizes and shapes, patternswlthm
geographical ranges (homerange,territory,habitat
selectlon), and limits to geographical distributions.
2 INTRODUCTION:
STUDYING
BIOGEOGRAPHY
Second, it examinesdispersal and range change,
considering how organisms move (agents of dispersal,
dispersal abilities, dispersal routes),
dispersal in
action, and dispersal In the past. Third, it explores
the splitting of geographical ranges (vicariance), dis-
cussing the effects of contlnental breakup on varlous
groups of organism (Triassic reptiles, Cenozolc land
mammals, Pangaean plants, large and flightless birds,
Greater Antillean insectivores). Fourth, I t looks at the
effects of continental fusion on life (faunal mixing and
the Great American Interchange). Fifth, it discusses
the ways In which humans a d and abet the spread of
organisms, using examples from the anlmal kingdom
(the American minkandmuntjac deer In Britain,
Introducedpredatorson Paclfic islands) andfungal
kingdom (the chestnut blight in the eastern Unlted
States).
Chapter 4 tackles populatlons. I t covers three
areas. First, it describes thedemography of single
populations,explainingthenature of population
growth (exponential, logistic, populatlon Irruptions,
population crashes, andchaotlcchange),ageand
sex structure (lifetables, survivorship curves, and
cohort-survival models), and metapopulations (loose,
tight, extinctlon-and-colonnation, andmainland-
Island). Second, I t outlines population survlval strate-
gles,
examining
opportunists
and
competitors
(v-strategists andK-strategists);competitors, stress-
tolerators,and ruderals; andmigrationstrategies
(fugitive,opportunist,andequilibrium).Thud, It
studies populatlon exploltatlon and control, lookmg
at overexploited populations (passenger plgeon,
auks,northern fur seal, and Americanbison), con-
trolledpopulations (Swedish beavers, NewZealand
goats, and Marion island cats), and the role of meta-
population theory in conservation (northern spotted
owl, common wallaroo or euro, Leadbeater’s possum).
Chapter 5 surveys Interacting populatlons. Itdevel-
ops five areas. First, I t looks at ways of livlng together
- protocooperatlon (anlmal-anlmal, plant-plant, and
animal-plant protocooperatlon, mimicry), mutual-
ism,andcommensalism(cattleegretsandcattle,
mldgeandmosquito larvae in pitcher-plant pools,
mynas andklng-crows, rlver ottersand beavers).
Second, i t looks at ways of staying apart, discusslng
types of competitlon (competitiveexclusion, scramble
competition,contestcompetltlon)andmechanisms
for avolding competltlon(resource partitioning, char-
acterdisplacement,spatlalcomplexity).Third, it
examines herbivory, looking at plant eaters (kinds of
herbivore and how plants defend themselves against
them) and plant-herbivore interactions (seed preda-
tionandgrazing).Fourth, it examinescarnlvory,
describing some aspects of flesh-eating (carnivorous
specializations, prey switching, prey selection, c a m -
vore communities),interactions between predators
and their prey (predator-prey cycles, chaos In Finnish
weasel and vole populations), and geographical effects
(laboratory and mathematlcalexperimentswlth
spatlally heterogeneous environments). Fifth, it shows
how life is pitted against life to control populations,
considertng biological control (prickly-pear cactus tn
Australia, false ragweed in Australia,agricultural
pests In the Mediterranean region), genetlc control,
and integrated pest management.
Chapter 6 delves Into communltles. It covers four
toplcs. First, I t describes the nature of communitles
and ecosystems, looking at a local ecosystem - the
NorthawGreatWood,England - andtheglobal
ecosystem. Second, it discusses roles wlthin commu-
nrtles, looklngatcommunityproductlon(prlmary
producers, prlmaryproduction),communitycon-
sumption (consumers,decomposers, and detritivores),
and ecosystem turnover (biogeochemicals, bio-
geochemical cycles). Thlrd, I t considers food chains
and food webs, exploring types of food web (grazlng
food chalns, decomposer food chams, ecologlcal pyra-
mlds), keystone species (keystone predators, keystone
herbivores and omnivores, the effects of removlng
keystonespecies), and
contaminated food webs
(biological magnificatlon, thelong-rangetransport
of radioactive isotopes andpesticides).Fourth, it
examines biodiversity, problng specles-area relation-
ships (specles-area curves, habitat diversity and
area-alonehypotheses),diversity gradients and hot-
spots,and diverslty change(why diversity matters
and how it can be safeguarded).
Chapter 7 I S about community change.I t expounds
four topics. First, it explains the nature of equilib-
rlumcommunlties,lookingattheclimaticclimax,
 INTRODUCTION: STUDYING BIOGEOGRAPHY 3

balancedecosystems, succession models (facilitation species under pressure and communitiesunder

model, tolerance model, inhibition model, allogenic
and autogenic changes), primary succession (Glacier
Bay, Alaska, Krakatau Islands,Indonesia), and sec-
.ondary succession (abandoned fields in Minnesota,
ghost towns in the western Great Basin). Second, it
explains the nature of disequilibrium communities,
discussing multidirectlonal succession (Hawaiian
montane ramforest, Glacier Bay again,Krakatau
Islands again)
and
communityImpermanence
(no-modern-analogue communities, disharmonious
communities, chaotic communities). Third,it tackles
land-cover transformation and its effect on communi-
ties, focusing on habitat fragmentation (the skipper
butterfly in Britain, the malleefowl in Australia, the
reticulated velvet gecko in Australia)andthe loss
of wetlands (bottomland forest in the SanteeRiver
floodplain in Georgia, coastal ecosystems in southern
Louisiana). Fourth, it predicts the possible shape of
communities in the twenty-first century, looking at
pressure (prairle wetlands in North America, mires in
the Prince Edward Islands, blome and ecotone shifts,
various examples of forest change induced by global
warming, lessons from ecologlcal simulations of
community change).
Chapter 8 turnstothehumandimension of
biogeography and ecology. It looks Into three areas.
First, it considers biorights, askmg if organisms have
rights(culling,thewildlifetrade, zoos) and if all
Nature has rights. Second, I t discusses attitudes
towards Nature, looking at brands of environment-
alism (technocentric, ecocentric), the alignments of
ecologists, and ecosystem management.Third, it
elucidatesthe
connection between
conservation
practiceand biogeographical and ecologlcal theory,
discussing envlronmental exploitation and enduring
equilibrium, balancedecosystems, evolutionary dis-
equilibrium, the edge of chaos, and the biodiversity
bandwagon.
 1

WHAT IS BIOGEOGRAPHY?
Biogeographers study the geography, ecology, a n d evolzrtion of living things. This chapter
covers:

ecology - environmental constraints on living

history and geography - time and space constraints on living

Biogeographers address a mdeadingly simple ques- teristlc life history, reproduction rate,behaviour,
twn: why do organisms live where they do? Why is means of dispersion, and so on. These traits affect a
the speckledrangeland grasshopper confined to population's response totheenvironment In whlch
short-grassprairie and forest or brushland clearings I t lives. The second idea concerns thls biological

containlng small patches of bare ground? Why does response to the environment and is the subject of
thering ouzel live in Norway, Sweden, theBrltlsh ecologlcal blogeography.Apopulation responds to
Isles, andmountalnousparts of centralEurope, its physical surroundings (abiotic environment) and
Turkey, and south-west Asla, but not In theinter- its
livlng
surroundings (biotic environment).
veningreglons?Whydotaplrs liveonly inSouth Factors In theabioticenvironmentinclude such
America andSouth-eastAsla?Whydothe nestor physlcal factors as temperature, light, soil, geology,
parrots - the kea and the kaka - live only in New topography, fire, water, water and alr currents; and
Zealand?Why do pouched mammals(marsupials) such chemicalfactors as oxygen levels, saltconcen-
live in Australiaandthe Americas, butnot in trations, the presence of toxms, and acidity. Factors
Europe, Asla, Afrlca, o r Antarctica? Why do differ- in the blotlc envlronment include competing species,
ent reglons carry distmct assemblages of animals and parasites, diseases, predators, and humans. In short,
plants? each species can toleratea range of environmental
Two groups of reasons are glven in answer to such factors. It can only live where these factors lie within
questmns as these - ecologlcal reasons and hlstorlcal- Its tolerance limlts.
cum-geographlcal reasons.
Speckled rangeland grasshopper
ECOLOGY This insect (Arphia conspwsa) ranges from Alaska and
northern Canada to northern Mexico, and from
Ecologicalexplanations for thedistributlon of California to the Great Plains. It is found at less than
organisms involve several interrelated Ideas. First 1,000 m elevation In the northern part of Its range
I S the idea of populations, whlch is the subject of andupto 4,000 m in thesouthernpart. Withm
analytical biogeography. Each specles has a charac- this extensive latitudinal andaltltudinalrange,its

distributionpattern is very patchy,owingto Its
decided preference for very specific habitats (e.g.
Schennum and Willey 1979). It requlres short-grass
pralrie, or forest and brushland openings, peppered
wlth small pockets of bare ground. Narrow-leaved
grasses provldethe grasshopper’s food source. The
barepatchesareneeded for it to perform courtship
rituals.These ecological and behavioural needs are
not met by dense forest, tall grass meadows, or dry
scrubland. Roadsidemeadows and old logged areas
are suitable and are slowly being colonized. Moder-
atelygrazedpasturesare also suitableandsupport
large populations.
Even withln suitable habitat, thegrasshopper’s dis-
tribution is limlted by Its low vagility (the ease with
which it can spread). Thls is the result of complex
social behaviour, rather than an inability to fly well.
Femalesare fairly sedentary,at least inmountain
areas, while males makemainlyshort,spontaneous
flights withln a limited area. The two sexes together
form tightly knit population clusters within areas of
suitablehabitat.Theclusters areheld together by
visual and acoustic communication displays.
Ring ouzel
The biogeography of most specles may be explained
by a mix of ecology and history. The ring ouzel or
‘mountain blackblrd’, whlch goes by the undignified
scientific name of Turdus torqrratus (Box l . l ) , lives in
the cool temperate climatic zone, and In the alpine
equivalent to the cool temperate zone on mountains
(Figure 1 . 1 ) . It likes cold climates. During the last
ice age, the heart of Its range was probably the Alps
and Balkans. From here, it spread outwardsinto
much of Europe, which was then colder than now.
With climatic warming durlng the last 10,000 years,
the rmg ouzel has left much of its former range and
survlves only in places that are still relativelycold
because of theirhlghlatitude or altltude. Even
though i t likes cold conditions, most ring ouzels
migrate to less severe climates during winter. The
north European populations move totheMediter-
ranean while the alpine populations move to lower
altitudes.
WHAT IS BIOGEOGRAPHY? 5
HISTORY A N D G E O G R A P H Y
Historical-cum-geographical explanations for the
distribution of organisms lnvolve two basic Ideas,
both of whlch are the subject of historical biogeog-
raphy. The first idea concerns centres-of-origin and
dispersal from one place to another. It argues that
species originate in a particular place and then spread
to other parts of the globe, if they should be able
and willing to do so. The second idea considers the
Importance of geologlcal and climatic changes split-
ting a single population into two or moreisolated
groups. This idea is known as vicariance biogeog-
raphy. These two baslc biogeographical processes are
seen in the following case studies.
Tapirs
The tapirs are close relatives of the horses and rhi-
noceroses. They form a family - the Tapiridae. There
are four living specles, one of which dwells in South-
east Asia andthree In CentralandSouth America
(Plate 1 .I). Their present distribution IS thus broken
and poses aproblem for biogeographers. Howdo
such closely related species come to live in geograph-
ically distant parts of the world? Findsof fossil tapirs
helpto answer this puzzle. Members of thetapir
family were once far more widely distributed than at
present (Figure 1.2). They are known to have lived in
North America and Eurasia. The oldest fossils come
from Europe. A logical
conclusion I S that
the
tapirs evolved in Europe, which was their centre of
origin, and then dispersed east and west. The tapirs
thatwent north-east reached North America and
South America. The tapirs thatchose a south-easterly
dispersal route moved intoSouth-east Asia. Subse-
quently,probablyowingtoclimaticchange,the
tapirs in North Amerlca and the Eurasian homeland
wentextinct.The survivors atthetropicaledges
of thedistributlon spawned the present species.
This explanation is plausible, though it is not water-
tight - it IS always possible that somebodywill
dig up even older tapir remainsfrom somewhere else.
Historical biogeographers are dogged by the incom-
pleteness of the fossil record, which means that they
can never be fully confident about any hypothesis.
6 WHAT I S BIOGEOGRAPHY?

i

Figure 1.1 The breeding distribution of the ring ouzel (Turdus twquatus).
Source: Map after Cramp (1988);picture from Saunders (1889)
Nestor parrots
The nestor parrots (Nestonnae) are endemic to New
Zealand.Thereare two specles - the kaka (Nestor
merzdionalis) and the kea (Nestor notubilis) (Plate 1.2).
They are closely related and are probably descended
from‘proto-kaka’
a that reached New Zealand
during the Tertiary p e r l o d . Then, New Zealand was
a single, forest-covered island. The proto-kaka
became adapted to forest life. Late in the Tertiary
period, the north and south parts of New Zealand
split. North Island remained forested and the proto-
kakastherecontinuedtosurvive as forestparrots,
feeding exclusively on vegetable matter and nesting
infree hollows. Theyeventuallyevolvedinto the
modern kakas. South Island gradually lost its forests
because mountains grew and climate changed. The
WHAT I S BIOGEOGRAPHY? 7
proto-kakas living on South Island adjusted to these
changes by becoming ‘mountain parrots’, depending
on alpine shrubs, insects, and even carrion for food.
They forsooktrees as breedingsitesandturnedto
rock fissures. The changes In the South Island proto-
kakas were so far-kaching that they became a new
species - the kea. After the Ice Age, climatic amelio-
ration promoted some reforestation of South Island.
The kakas dispersed across the Cook Strait and colo-
nized South Island. Interaction between North and
South Island kaka populations 1s difficult across the
26 km of ocean. Inconsequence, the South Island
kakas have become a subspecies. The kaka and the
kea are now incapable of interbreeding and theycon-
tmue tolive sideby side on South Island. The kea has
never colonized North Island, probablybecause there
islittlesuitablehabitatthere.Thebiogeography
 9
” rr
a d

i
ii

Plate 1.1 Central American or k i d ‘ s tapir (Sapirus h i d ) , Belize

Photograph by Pat Morris

of the nestorparrotsthusinvolvesadaptationto The classic explanation of marsupla1 distribution

changmgenvironmentalconditions,dispersal,and
vicariance events.
Marsupials
The pouched mammals or marsupials are now found
inAustralia,SouthAmerica,andNorthAmerica.
Fossil forms are known from Eurasia, North Africa,
and Antarctica. They are commonly assumed to have
evolved in North America, where the oldest known
marsupial fossils are found, from an ancestor that also
sitedtheplacentalmammals.Themarsupialand
placentalmammalssplitabout 130 million years
ago, early in the Cretaceous period. There are several
rivalexplanationsfor thecurrentdistributlon of
marsupials (L.G . Marshall 1980).
was proposed before continental drift was accepted
andassumedstatlonarycontments. Itarguedthat
marsupialsdispersed
from
Cretaceous
a North
American homeland to other continents. Some time
inthe LateCretaceousperiod,marsupialshopped
acrossislandslinkingNorthandSouthAmerica.
During the Eocene epoch, they moved into Asia and
Europe acrossaland bridgespanningtheBerlng
Sea betweenAlaskaandSiberia.Fromtherethey
spreadintoEuropeand,usingIndonesia as an
embarkation point, into Australia. Several variations
onthls‘centre of orlgln followedbydispersal’
hypothesisplayed outonastationarylandsurface
were forthcoming. The variations involved different
centres of origin (South America or Antarctica) and
different dispersal routes.
 9

Figure 1.2 Tap~n:their origm, spread, and present distribution.

Source: After Rodriguez de la Fuente (1975)

Plate 1.2 Nestor parrots (a) Kaka (Nestor rnm'dionulk) (b)Kea (Nestor notubilk)
Photographs by Pat Morris
10 WHAT IS BIOGEOGRAPHY?
As soon as it was accepted that the continents do
drift, revised explanations of the marsupialhistory
were suggested. Some of these new hypotheses still
Invoked centre-of-origin and dispersal. They were
similartothe hypothesisdeveloped for stationary
continents, but they did not need to invoke fanciful
land bridges betweenwidelyseparated continents.
Other hypotheses laid emphasls on the fragmentatlon
of Pangaea, the Triassic supercontinent, and stressed
vicarlanceevents ratherthan dispersaloverpre-
exlsting barriers.
It now seemslikely that, even if the first mar-
supials did appear in Mesozoic North America (and
that IS far from certain), they quickly became wldely
distributed over the connected land masses of South
Amerlca, Antarctica, and Australia. The breakup of
Pangaea,which started inearnest duringtheMid
Jurasslc period, Isolated the Mesozolc marsupials on
South America andAustraliaand these twomam
branches then evolved independently. The American
marsupials reached Europe, North Africa, and Asia in
Palaeocene and Eocene tlmes, but by the end of the
Miocene epoch, North American and European mar-
supials were extinct.South Americanmarsupials
invaded North Americain the Pleistocene epoch.
Current explanatlons of marsupial biogeography thus
call on dispersal and vicariance events.
Biogeographical regions
Different places house different kinds of animals and
plants.This became apparent as the world was
explored. In 1628, in his The Anatomy of Melancholy,
Robert Burton wrote:
Whydoth Afrlcabreed so manyvenomousbeasts,
Irelandnone?Athensowls,Cretenone?Whyhath
Daulis and Thebes no swallows (so Pausanlas informeth
us) as well as the rest of Greece, Ithaca no hares, Pontus
[no]
asses, Scythia[no]wlne?Whencecomesthis
varletyofcomplexlons,colours,plants,blrds,beasts,
metals, peculiar to almost every place?
(Burton 18% edn: ~ 0 1 11,
. 50-1)
By the nineteenth century, it was abundantly clear
that the land surface could be divldedinto several
large biogeographical regions, each of which sup-
ports a distmct set of animals and a distinct set of
plants.Usingblrddistributions,Philip L. Sclater
(1858) recognized two basic divisions - theOld
WorldandtheNewWorld.TheOldWorld he
divided into Europe and northern Asia, Africa south
of the Sahara, India and southern Asia, and Australia
and New Guinea. The New World he divided into
North Amerlca and South America. Sclater’s system
was adopted by Alfred Russel Wallace (1876),
with minor amendments, to provide along-lasting
nomenclaturethat survivestoday as the Sclater-
Wallace scheme (Figure 1.3a). Six regions are recog-
nlzed - Nearctlc, Neotroplcal, Palaearctic, Ethioplan,
Oriental, and Australian. Together, the Nearctlc and
PalaearcticformNeogaea (theNewWorld),while
other regionsform Palaeogaea (TheOldWorld).
Wallace also recognized subregions, four per region
(Table l . l ) , whlch correspond largely to established
plant regions.
Modernmethods of numerical classification pro-
duce similar regions to the Sclater-Wallace scheme,
butthere aredifferences. Figure1.3b showsfaunal
regions of the world based on mammal distributlons
(C. H. Smith 1983). There are four regions - Hol-
arctic, LatmAmerican,Afro-Tethyan,and Island
- and ten subregions. Each subregion is as unlque as
it can be compared with all other subregions.
SUMMARY
Thedistribution of organisms is determined by
ecology, history, and geography. Most distributions
resultfrom a combination of all three factors. Bio-
logical and geological evolution have acted together
toproducethe blogeographicalregions andsub-
regions seen today.
E S S A YQ U E S T I O N S
1 Describethecharacteristicanimalsin
Wallace’s biogeographical regions.
2 Describethechiefbotanicalregionsof
the world.
 11

Nearctic
0Neotropical
a Palaearctic

Ethiopian

Oriental
Australian

Island

Figure 1.3 (a) TheSclaterandWallaceclassificatlon of faunal regions.(b)Anumerlcalclassificatlon of mammal

distributlons showlng four maln reglons and ren subregions.
Source: After C. H. Smith (1983)
12 W H A T IS BIOGEOGRAPHY?

Table I . I Biogeographical regions and subregions,as defined by Alfred Russel Wallace

Region Subregion

Palaeogaea (Old World)

Palaearctic North Europe
Mediterranean
Siberia
Manchuria (or Japan)
Ethiopian East Africa
West Africa
South Africa
Madagascar
Oriental Hindustan (or central India)
Ceylon (Sri Lanka)
Indochina (or Himalayas)

Australian IndoMalara
Austro-Ma aya
Australia
Polynesia
New Zealand

Neogaea (New World)

Neotropical Chile
Brazil
Mexico
Antilles
Nearctic California
Rocky Mountains
Alleghenies
Canada

Source: After Wallace (1 876)

F U R T H E R READING

COX,C. B. and Moore, P. D. (1993) Biogeography: A n

Ecological and Evolntronav Approarb, 5th edn, Oxford:
Blackwell.

George, W. (1962) Animal Geography, London:

Heinemann.

Tivy, J. (1992) Biogeography: A Study of Plants in the

Ecosphere, 3rd edn, Edinburgh: Oliver & Boyd.
 2
LIFE AND THE ENVIRONMENT
Life i s adapted to nearly all Earth surface environments. This chapter covers:
placestolive
climatic constraints on living
other physical constraints on living
ways of living
LIVING S P A C E : H A B I T A T S A N D
ENVIRONMENTS
Individuals, species, andpopulatlons,bothmarine
and terrestrial, tend to live in particular places. These
places arecalled habitats. Each habitat I S charac-
terlzed by a speclfic set of environmental conditlons
- radiation and light, temperature, molsture, wind,
fire frequency andintenslty,gravity,salinity,cur-
rents,topography, soil, substrate,geomorphology,
human disturbance, and so forth.
A place to live: habitats
Habitats come in all shapes and sues, occupying the
full sweep of geographical scales. They rangefrom
small
(microhabltats),
through
medium (meso-
habitats) and large (macrohabltats), to very large (mega-
habitats). Microhabitatsare a few square centimetres
toa few squaremetres in area (Table2.1).They
Include leaves, the soil, lake bottoms, sandy beaches,
talus slopes,walls, river banks,andpaths.Meso-
habitats have areas up to about 10,000 km’; that is,
a 100 x 100kilometresquare, which is about the
size of Cheshire, England. Each main mesohabltat is
Influenced by the same reglonal climate, by similar
features of geomorphology and soils, and by a similar
set of disturbancereglmes.Deciduouswoodland,
caves, and streams are examples. Macrohabitats have
areas up to about 1,000,000 km‘, which IS about the
size of Ireland. Megahabitats are regions more than
1,000,000 km’ in extent.They Include continents
and the entlre land surface of the Earth.
Landscape ecologlsts, who have an express interest
In the geographlcal dimension of ecosystems, recog-
nize three levels of ‘habltat’ - region, landscape, and
landscape element.These correspond to large-scale,
medium-scale, and small-scale habitats. Some land-
scape ecologists are relaxing their Interpretation of a
landscape to Include smaller and larger scales - they
have come to realize that abeetle’s view or a bird’sview
of the landscape I S very different from a human’sview.
Landscape elements
Landscape elements are similartomicrohabitats,
but a little larger. They are fairly uniform pieces of
land, no smallerthanabout 10 m,that form the
building blocks of landscapes and regions. They are
also calledecotopes, biotopes,
geotopes, facies,
sites,tesserae,landscape unlts, landscapecells, and
landscape prisms. These terms are roughly equivalent
to landscape element,but have theirown speclal
meanlngs (see Forman 1995; Huggett 1995).
14 L I F E A N D THE ENVIRONMENT

Table 2. I Habitat scales

Approximate
Scaleo area Terminology applied to landscape units at same scaleb
/km21
Fenneman linton Whittlesey
(1916) [ 1949) [ I 954)
Microhabitat <1 - Site -
(small)
Mesohabitat 1-10 - - -
(medium) 10-1 00 - stow Locality
1 00-1,000 District Tract District
1,000-1 0,000 Section Section -
Macrohabitat 10,000-1 00,000 Province Province Province
(large) 100,000-1,000,000 Major division Major division Realm
Megahabitat > 1,000,000 - Continent -
(very large)

Note: a ThesedivisionsfollowDelcourtandDelcourt ( 1 988)

The range of areas associated with these regional landscape units is meantas a rough-and-ready
guideratherthanprecise limits

Landscape elements are made of individual trees,
shrubs, herbs, and small buildings. There are three
basic kinds of landscape element - patches, corridors,
and background matrixes:
1 Patches are fairly uniform(homogeneous) areas
that differ from their surroundings. Woods, fields,
ponds, rock outcrops, and houses are all patches.
2 Corridors are strips of landthat differfrom
the land to either side. They may interconnect to
form networks. Roads, hedgerows, and rivers are
corridors.
3 Background matrixes are thebackground eco-
systems or land-usetypes in which patchesand
corridorsare set. Examplesare deciduous forest
and areas of arable cultivation.
Landscape elements include the results of human toil
- roads, railways, canals, houses, and so on. Such fea-
tures dominate the landscapeinmany parts of the
world and form a kind of ‘designer mosaic’. Designed
patches includeurban areas, urban andsuburban
parks and gardens (greenspaces), fields, cleared land,
and reservoirs. Designed corridorsinclude hedge-
rows, roads and railways, canals, dikes, bridle paths,
and footpaths. There is also a variety of undesigned
patches - waste tips, derelict land, spoil heaps, and
so on.
Landscapes
Landscape elements combine to form landscapes. A
landscape is a mosaic, an assortment of patches and
corridors setin amatrix, no biggerthanabout
10,000 km’. It is ‘a heterogeneousland area com-
posed of a cluster of interacting ecosystems that is
repeatedin similarformthroughout’(Formanand
Godron 1986: 11). By way of example, the recurring
cluster of interacting ecosystems that feature in the
landscape around the author’s home, in the foothills
of the Pennines, includes woodland, field, hedgerow,
pond,brook, canal,roadside, path, quarry, mine
tip, disused mlningincline, disusedrailway,farm
building, and residential plot.

The bare necessities: habitat
requirements
It I S probably true to say that no two specles have
exactly the same living requirements. There are two
extreme cases - fussy specles or habitat specialists
and unfussy species or habltat generalists - and all
grades of 'fussmess' between.
Habitat specialists
Habitat specialists have very precise living require-
ments.InsouthernEngland,the red ant, Myrrnira
sabuleti, needs dryheathlandwitha warm south-
faclngaspect thatcontains more than 50 percent
grass species, and that has been disturbed within the
previous five years (N. R. Webb and Thomas 1994).
Other species are less pernicketyandthrive over a
wider range of environmental conditions. The three-
toed woodpecker (Piroides triducryh)lives in a broad
swath of cool temperate forest encircling the North-
ern Hemisphere. Races of the common jay ( G a w u h s
glundurirts) occupy a belt of oak and mixed deciduous
woodland stretching from Britain to Japan.
Habitat generalists
A few species manage to eke out a living in a great
array of environments.Thehuman species (Homo
sapiens) is thechampionhabitatgeneralist - the
planet Earth is the human habitat. In the plant king-
dom,the broad-leaved plantain (Plantago mujor),
typically a species of grassland habitats, is found
almost everywhereexcept Antarcticaandthedry
parts of North Africa and the Middle East. In the
Brlttsh Isles, it seems indifferent to climate and soil
LIFE A N D THE ENVIRONMENT 15
Life's limits: ecological tolerance
Organisms live In virtually all environments, from
the hottest to the coldest, the wettest to the drlest,
the mostacldic tothe most alkaline.Understand-
ably,humanstendtothlnk of their'comfortable'
envlronment as the norm. But moderate conditlons
are anathema to the mlcro-organisms that love con-
ditions fatal to other creatures. These are the extre-
mophiles(Madiganand Marrs 1997). An example
is high-pressure-loving microbes (barophiles)that
flourlsh In deep-sea environments and are adapted to
life athigh pressures (Bartlett 1992). Many other
organisms are adapted to conditions that, by white
western human standards, areharsh, though not so
extreme as the conditions favoured by the extremo-
philes.Examplesare hot deserts and Arctic and
alpine regions.
Limiting factors
A limiting factor is an environmental factor that
slows down population growth. The term was first
suggested by Justus von Liebig (1840), aGerman
agricultural chemist. Liebig noticed that the growth
of a field crop is limited by whichever nutrient
happens to be in short supply. A field of wheat may
have ample phosphorus to yield well, but if another
nutrient, say nitrogen, should be lacking, then the
yield will be reduced. No matter how muchextra
phosphorus I S applied in fertilizer, the lack of nitro-
gen will limlt wheatyield. Only by makinggood
thenitrogenshortagecould yields be improved.
These observations led to Liebig to establish a 'law
oftheminimum':theproductivlty,growth,and
reproduction of organisms will be constrained if one
16 LIFE AND THE ENVIRONMENT

or more environmental factors lies below its limiting Each species (or race) has a characteristic toler-
lwel. ance range (Figure 2.2). Stenoecious species have a
Later, ecologlsts established a ‘law of the maxi- widetolerance;euryoecious species have anarrow
mum’.This law applies where population growth is tolerance. All species, regardless of theirtolerance
curtailed by anenvironmentalfactorexceedingan range, may be adapted to thelow end (oligotyplc), t o
upperlimiting level. Inawheat field, toomuch the middle (mesotypic), or to the high end (poly-
phosphorus is as harmhl as too little - there is an typic) of an environmental gradient. Take the exam-
upper limn to nutrient levels that plants can tolerate.ple of photosynthesisinplants.Plantsadaptedto
cool temperatures (oligotherms) have photosynthetic
Tolerance range optima at about 10°C and cease t o photosynthesize
above 25OC. Temperate-zoneplants(mesotherms)
For wery environmental factor (such as temperature have optima between 15OC and 30°C. Tropical plants
and moisture) there is a lower limit below which a (polytherms) may have optlma as high as 4OOC.
species cannot live, an optimum range In which it Interestingly,theseoptimaarenot‘hardand fast’.
thrives, and an upper limit above which it cannot live Cold-adaptedplantsareabletoshifttheirphoto-
(Figure 2.1). The upper and lower bounds define the synthetic optima towards higher temperatures when
tolerance range of a species for a partlcular envlron- they are grown under warmer conditlons.
mental factor. Thebounds vary fromspecles to
species. A species will prosper within Its optimum
range of tolerance; survive but show signsof physio- Ecological valency
logical stress near its tolerance limits; and not survlveTolerance may be wide or narrow and the optimum
outside Its tolerance range (Shelford 1911). Stress is may be at low, middle, or high posltions along an
a widely used but troublesome idea in ecology. It environmental gradient. When combmed, these con-
may be defined as ‘external constraints limltlng the tlngencies produce six grades of ecological valency
rates of resource acquisition, growth or reproduction (Figure 2.2). The glacial flea (Isotoma saltans) has a
of organlsms’ (Grime 1989). narrow temperature tolerance and likes it cold. It IS

I range Tolerance

Organisms
absent

Figure 2.1 Tolerance range and limits.

Source: Developed from Shelford (1911)
 LIFE A N D THE ENVIRONMENT 17

Examples:
100
a. Oligostenotherm - Glacial flea
(kotorna saltans)
b. Mesostenorheob - Barbel
(Barbus fluviatilis)
c. Polystenoxyblont - Midge
(Liponeura cinerascens)
Tolerance
(per cent) 50 d. Oligoeurybiont - Chironomid
(Chironomus plumosus)
e. Mesoeuryrheob - Freshwater limpet
(Ancylus fluvratilis)
f. Polyeuryhalob - Jelly-fish
(Aurelia aurita)

I Environmental
factor
Oligo- I Meso- I Poly-

Oxygen (mg/l) Oxybiont

Temperature ("C 10-20 Therm
Salinity (V,J 0-30 30-35 Halob
Current (mk) 0-0.05 0.05-0.5 >0.5 Rheob

F i p r e 2.2 Ecological valency, showing the amplitude and position of the optimum
Source: After Illies (1974)

an oligostenotherm. The midge Lzponewa rinwasl-ens
has a narrow oxygen-level tolerance at the high end
of the oxygen-level gradient. It is a polystenoxybiont.
Other examples are shown on Figure 2.2.
WARM AND WET: CLIMATIC
FACTORS
Flower power: radiation and light
The Sun I S theprlmary source of radiation for the
Earth.Itemltselectromagnetic radiation across a
broad spectrum, from very short wavelengths to long
wavelengths (Box 2.1). The visible portion (sunlight)
is the effectlve bit for photosynthesis.It is also
significant In heatingtheenvironment. Long-wave
(infrared)radiationemitted by theEarth is locally
Important around volcanoes, in geothermal springs,
andinhydrothermalvents in the deep-sea floor.
Theseinternal sources of energy are tapped by
unusual organisms, Including the thermophiles and
hyperthermophiles that like I t very hot (p. 19).
Three aspects of solar radiation influence photo-
synthesis - the Intensity, the quality, and the photo-
period or duration. The Intensity of solar radiation is
the amount that falls on a given area in a unit of
tlme. Calorlesper squarecentlrnetreperminute
(cal/cm'/ min) wereonce popular units, but Watts
per square metre (W/m*) or kiloJoulesperhectare
(kJ/ha)aremetricalternatives.The average annual
solar radiation on a horizontal ground surface ranges
from about 800 kJ/ha over subtropical deserts to less
than 300 kJ/ha in polarregions. Equatorial regions
receive less radiation than the subtropicsbecause they
are cloudier. A value of 700 kJ/ha is typical.
Thequality of solar radiation is itswavelength
composition. This vanes from place to place depend-
Ing on the composltlon of the atmosphere, different
components of whlch filter out different parts of the
electromagnetic spectrum. In the tropics, about twice
as much ultraviolet light reaches the ground above
2,500 m than at sea level. Indeed, ultravlolet light is
stronger in all mountains - hence incautious humans
may unexpectedly suffer sunburn at ski resorts.
Photoperiod IS seasonal variations In the length
of day andnlght.This is immenselyimportant
ecologically because day-length, or more usually
18 LIFE AND THE ENVIRONMENT

Box 2.1
T H EE L E C T R O M A G N E T I CS P E C T R U M lengthsintherange 0.4 to 0.8 pm. This is the
E M I T T E D BY T H E S U N portion of the electromagneticspectrumhumans
can see. Infrared radiation has wavelengths longer
Electromagnetic radiation pours out of the Sun at than 0.8 pm. It grades into radio frequencies with
the speed of light.Extremeultravioletradiation millimetretometrewavelengths.TheSunemits
withwavelengthsintherange 30 to120 nano- mostintenselynear5 prn, which is in the green
metres(nm)occupiesthe very shortend of the band of the visible light. This fact might help t c
spectrum. Ultraviolet light extends to wavelengths accountforplantsbeinggreen - theyreflect thc
of 0.4 micrometres (pm). Visible light has wave- most intense band of sunlight.

night-length, stimulates the timrng of daily and sea- Arctlc and alpine animals and plants also have to
sonal rhythms (breeding, migration, flowering, and so cope with limited solar energy. Herblvores gear their
on) in many organisms. Short-day plants flower when behaviour to making the most of the short summer.
day-length IS below a critical level. The cocklebur Belding ground squlrrels (Spermophilus beldingi),
which
(Xanthzumstncmarzum), a widespread weed in many live at high elevations In the western United States,
parts of the world, flowers In spring when, as days are actwe for four or five summer months, and they
become longer, a critlcal night-lengthIS reached (Ray must eat enough durlng that time to survive the win-
and Alexander 1966). Long-day plants flower when ter on stored fat (Morhardt and Gates 1974). To do
day-length ISabove a critical level. The strawberry tree thls, their body temperature fluctuatesby 3 4 ° C (to a
(Arbutus unedo) flowers In the autumn as the night- hlgh of 40°C)so that valuable energyIS not wasted In
length increases. InitsMediterraneanhome,this keeping body temperature constant. Should they need
means that Its flowers are ready for pollination when to cool down, they go into a burrow or else adopt a
such long-tongued Insects as bees are plentihl. Day- posture that lessens exposure to sunlight. A constant
neutralplants flower afteraperiod of vegetative breeze cools them durlng the hottest part of the day.
growth, irrespectlve of the photoperiod.
In the high Arctlc, plant growth is telescoped Into
Some like it hot: temperature
a brief few months of warmthandlight.Positive
heliotropism (growing towards the Sun) IS one way Broadly speaklng,
average
annual
temperatures
that plants can cope with limited light. 1sItcommon are highest at the equator and lowest at the poles.
in Arctlc and alpine flowers. The flowers of the Arctic Temperatures also decrease with lncreaslng elevation.
avens (Dtyas integr$oIia) and the
Arctic
poppy The average annual temperature range IS an impor-
(Papaw radicatum) track the Sun, turning at about tantecologlcalfactor.It IS hlghestdeep In high-
15" of arc per hour (Kevan 1975; see also Corbett et latltude contrnental interlors and lowest over oceans,
al. 1992). Thelr corollas reflect radiation onto their especiallytropicaloceans.Innorth-eastSiberia,an
reproductive parts. The flowers of the alpine snow annual temperature rangeof 60°C IS not uncommon,
buttercup (Ranunculus adoneus) track the Sun's move- whereas the range over equatorial oceans IS less than
mentfrom
early
morninguntil
mld-afternoon about 3°C. Land lying adjacent to oceans, especially
(Stanton and Galen 1989).Buttercup flowers aligned land on the western seaboard of conrlnents, has an
parallel to the Sun's rays reach mean internal tem- annualtemperaturerangearoundthe11°Cmark.
peratures several degrees Celsius above ambient air These large differences in annual temperature range
temperature. Internal flower temperature is slgnifi- reflect differences incontinentality (or oceanicity)-
cantly reduced as a flower's angle of devlatlon from the wlnter temperatures of places near oceans will be
the Sun increases beyond 45". less cold.
 LIFE AND THE ENVIRONMENT 19

Manyaspectsoftemperature affect organlsms, Below 94°C I t finds I t too cold and stops growing!
Includingdaily,monthly,andannualextremeand Only In small areas thatareintenselyheated by
meantemperatures,andthe levelof temperature volcanic activity do high temperatures prevent life.
variability. Different aspects of temperature are rele- Cold-lovlng mlcrobes (psychrophiles) are common
vant to differentspeclesandcommonly vary wlth inAntarctic sea ice. Thesecommunitiesinclude
the time of year and the stage in an organlsm's life photosynthetic algae and diatoms, and a varlety of
cycle. Temperature may be limitlng at any stage of bacteria. Polaromonas zwcxolata, abacterium,grows
an organlsm's life cycle. It may affect survival, repro- best atabout 4"C, andstopsreproducingabove
ductlon, and the developmentof seedlings and young 12°C. Lichens can photosynthesize at -3O"C, provid-
animals. It may affect competitlon with other organ- ingthattheyarenot covered withsnow.The
isms and susceptibility to predation, parasitism, and reddish-colouredsnowalga, Ch1an1yrfomona.r nrvdfis,
disease whenthelimitsoftemperaturetolerance lives on ice and snow fields In thepolarand nival
are approached. Many flowering plants are especially zones, giving the landscape a pink tinge durmg the
sensitive to low temperaturesbetweengerminatlon summer months.
and seedling growth.
Animals a n d temperature
Microbes and temperature
Inmostanlmals,temperature is a critical limitmg
Heat-loving microbes (thermophiles) reproduce or factor. Vital metabolic processes are geared to work
grow readily In temperatures over 45°C. Hyperther- optimally withrn a narrow temperature band. Cold-
mophiles, such as Suljolobl/~~~ ' . j [ f o l ~ / ~ ( f ' l r prefer
l//~, blooded anlmals (poikilotherms) warm up and cool
temperaturesabove 80°C, andsomethriveabove down wlth environmenral temperature (Figure 2.321).
100°C. Themost resistanthyperthermophile clis- Theycan assist thewarming process alittle by
covered t o date is Pyrolobl/.r j i t m r i i . Thlsmicrobe taking advantage of sunny spots or warm rocks. Most
flourishesin the walls of'smokers'inthe deep-sea warm-blooded animals (horneotherms) maintain a
floor. Itmult~plies In temperatures up to 113°C. constant body temperature amidst varylng ambient

e 30

0 I I 1 I I
0 10 20 30 40 0 10 20 30 40
Environmental temperature ("C) Environmental temperature ("C)

Fipwe 2..i Temperature control i n poikilotherms and homeotherms

So~wrr:After Bartholomew (1 968)
20 L I F E A N D THE ENVIRONMENT
conditions(Figure2.3a).Theysimplyregulatethe
production and dissipation of heat. The terms 'cold-
blooded' and 'warm-blooded' are misleading because
the body temperature of some 'cold-blooded' animals
may rise above that of 'warm-blooded' animals.
Each homeothermlc species has acharacteristic
thermal neutral zone, a band of temperature within
which little energy is expended In heat regulatlon
(Figure2.3b)(Bartholomew1968).Smalladjust-
ments are made by fluffing or compressing fur, by
making local changesin the blood supply, or by
changmg position. The bottom end of the thermal
neutral zone I S bounded by a lower critical temper-
ature. Below thls temperature threshold, the body's
central heating system comes on fully. The colder i t
gets,the moreoxygen is needed toburn fuel for
heat. Animals living incold environments arewell
insulated - furandblubber can reduce the lower
33
Figure 2.4 Temperatures at an Esklmo dog's extremities ("C) with an ambient a ~ temperature
S u r m ~ :After Irving (1966)
criticaltemperatureconsiderably. An Arctic fox
(Alopex lagopus) clothed in its wmter fur rests com-
fortably at an ambient temperature of-50°C without
increasing itsresting rate of metabolism(Irving
1966). Below the lowercritical temperature,the
peripheral circulation shuts down toconserve energy.
An Eskimo dog may have a deep body temperature
of 38"C, the carpal area of the forelimb at 14"C, and
foot pads at O°C (Irving 1966) (Figure 2.4). Hollow
hair I S also useful for keeping warm. I t is found in the
American pronghorn (Antilocapra amwzcana),an even-
toed ungulate,and enables it to stay inopen and
windswept places at temperatures far below OOC. The
polar bear (Ursus nzarrfzmus) combines hollow hair, a
layer of blubber up to 11 cm thick, andblack skin to
produce a superb insulating machine. Each hair acts
like a fibre-opticcable, conducting warming ultra-
violet lighttotheheat-absorbing black skin.This
r of-30°C
 LIFE AND THE ENVIRONMENT 21

heating mechanism is so efficient that polar bears are

more likely to overheat than to chill down, which Bushy-tailed woodrat
2 (Neotoma cinerea)
partlyexplainstheirponderousness.Manyanimals Oregon
also have behavloural patterns deslgned to minimize
heat loss. Some roll into a ball, some seekshelter.
4

2L “L
Herds of deer or elk seek ridge tops or south-facing
Bushy-tailed woodrat
slopes. (Neotoma cinerea)
Above the upper critical temperature, animals
must lose heat to prevent their overheating. Anlmals
0 I
livinginhotenvironments canlosemuchheat.
al
Evaporationhelpsheat loss, but has anunwanted White-throated woodrat
VI
side-effect - preclous water IS lost. Small animals can 5
m
burrowtoavoldhightemperaturesattheground aJ
U
.c
surface. In the Arlzona desert, Unlted States, most 0
rodents burrow to a depth where hot or cold heat
stress is not met with. Large size is an advantage In
preventlng Overheatingbecause the surfaceareais
relatively greater than thebody volume. Many desert
mammalsareadaptedtohlghtemperatures.The 6
African rock hyrax (Heterobyrax&rei) has an upper White-throated woodrat
(Neotoma albigula)
critical temperature of 41°C. Camels (Camelus spp.), 4
oryx (Oryx spp.), common eland (Taurotragusoryx), and
2
gazelle (Gazella spp.)lettheirbodytemperatures
fluctuate considerably over a 24-hour period, falling
0
to about 35°C toward dawn and rlsing to over 40°C 33 37 41 45
durlng the lateafternoon. In an ambient temperature Ambient temperature (“C)
of 45°C sustalned for 12 hours under experlmental
conditions, anoryx’s temperatute rose above 45°C and Figure 2.5 Lethal amblent temperatures for fourpopula-
tlons of woodrats (Neotm) living in the western United
stayed there for 8 hours wlthout injuring the animal
States. The numbers of deaths, shown by the shaded areas,
(Taylor 1969). It has a specialized circulatory system are based on four-hour exposures.
that helps I t to survive such excessive overheating. Sourre: After Brown (1968)
Many mammal species are adapted to a limlted
range of environmentaltemperatures. Evenclosely transpiration. Plants vary enormously in their ability
related groups display significant differencesIn them to tolerate either heat or cold. There are five broad
ability to endure temperature extremes. The lethal categorles of coldtolerance(Table 2.2). Chilling-
ambient temperatures for four populatlons of wood- sensitiveplants,whicharemostlytroplcal,are
rats (Neotoma spp.) In the westernUnltedStates damaged by temperatures lower than10°C. Chilling-
showeddifferencesbetweenspecies,andbetween resistant(frost-sensitive)plants cansurvive at
populations of the same sp&ies livlng in different temperatures below 10°C, but are damaged whenice
states (Figure 2.5). formswithlntheirtissues.Frost-resistantplants
makephysiologicalchanges that enable themto
Plants and temperature survive temperatures as low as about -15°C. Frost-
tolerant plants survive by withdrawing water from
Temperature affects many processes in plants, including their cells,so preventing ice forming. The withdrawal
photosynthesis, respiration, growth, reproductlon, and of water also increases the concentratlon in sap and
22 LIFE AND THE ENVIRONMENT
Table 2.2 Temperature tolerance in plants
Temperature sensitivity Minimum temperature
life-form
("Cl
Chilling-sensitive >10
Chilling-resistant (frost sensitive) 0 to 10
Frost-resistant -15 to 10
Frost-tolerant -40 to -15
Cold-tolerant <- 40
Source: After Woodward ( 1 992)
protoplasm, which acts as a kind of antifreeze, and
lowersfreezing point. Temperatures down to about
-40°C can be tolerated in this way. Cold-tolerant
plants, which are mostlyneedle-leaved, can survive
almost any subzero temperature.
Cold
tolerance varies enormously at different
seasons in some species. Willowtwlgs (Sulix spp.)
collected in wlnter can survive freezing temperatures
below -150°C; the same twlgs in summer are killed
by a temperature of -5°C (Sakai 1970). Similarly, the
red-osler dogwood (Cornr/s stolonifera), a hardy shrub
from North America, could survive a laboratory test
at -196°C by midwinter when grown in Minnesota
(Weiser1970).Nonetheless,dogwoods native to
coastal regions wlth mild climates are often damaged
by early autumn frosts. Plants growlng on Mt
Kurodake, Hokkaldo Provlnce, Japan, are killed by
temperatures of -5°C to -7°C during the growmg
season. Inwlnter, most of the same plants survive
freezing to -30"C, and the willow ezo-mame-yanagi
(Sulix puuciflora), mosses, and lichens will withstand
a temperature of -70°C (Sakai andOtsuka1970).
Acclimatization or cold hardening accounts for these
differences. The coastal dogwoods did not acclimatlze
quickly enough. Timlng is important incold resis-
tance, butabsolute reslstance can be altered. Many
plants use the signal of short days in autumn as an
early warning system. The short days trigger meta-
bolic changes that stop the plant growing and pro-
duce resistance to cold. Many plant specles, especially
deciduous plants In temperate regions, need chilling
durlng winter if they are to grow well the followmg
Broad-leaved evergreen
Broad-leaved evergreen
Broad-leaved evergreen
Broad-leaved deciduous
Broad-leaved evergreen and
deciduous; boreal needleleaved
summer. Chilling requirementsare specific to species.
Theyare often necessary for budsto break out of
dormancy, a process called vernalization.
Many plants require a certain amount of 'warmth'
durmg the year. The total 'warmth' depends on the
growing season length and
the
growing season
temperature. These two factors are combined as 'day-
degree totals' (Woodward 1992). Day-degree totals
are theproduct of thegrowing season length(the
number of days for which the mean temperature I S
above standardtemperature, such as freezing pomt
or S"C), and the mean temperature for that period.
The Iceland purslane (Koenzxia rsluwdizu), atundra
annual, needs only 700 day-degrees to develop from
a germinating seed to a mature plant producingseeds
of its own. The small-leaved lime (Tiliu mrdutLz), a
declduous tree, needs 2,000 day-degrees to complete
its reproductive development (Pigott 1981). Trees in
tropical forests may need up to 10,000 day-degrees to
complete their reproductive development.
Excess~ve heat is as detrimentaltoplants as
excessive cold. Plants have evolved reslstance to heat
stress, thoughthe changesare not so marked as
reststance to cold stress (see Gates 1980, 1993: 69-
72). Different parts of plants acquire differingdegrees
of heatresistance,bur thepatternvanes between
species. In some specles the uppermost canopy leaves
are often the most heat reslstant; In other species, I t I S
the middle canopy leaves, or the leaves at the base
of the plant. Temperatures of about 44°C are usually
injurious
to evergreens andshrubsfrom
cold-
wlnter regions. Temperate-zone trees are damaged at
 LIFE AND THE ENVIRONMENT 23

50-55"C, tropical treesat 45-55"C. Damage Incurred heat and stress the plant. Similarly, for a plant to use
belowabout50°C can normally be repalred by the water for growth, energy must be obtainable. With-
plant;damageincurredabovethattemperature IS out an energy source, the water will run into the soil
mostoftenIrreversible.Exposure tlme to excess~ve orrun off unused. For theserrasons,temperature
heat is a crltlcal factor in plant survival, while exposure(as a measureofenergy)andmolstureare master
time to freezing temperatures is not. limiting factors that actintandem.Intropcal
areas, temperatures are always hlgh enough for plant
growthandprec~pltation I S thelimlting factor. In
Quenching thirst: moisture
cold environments,water is usuallyavailable for
Protoplasm,thelivingmatterofanlmalandplant plant growth for most of the year - low temperatures
cells, is about 90 per cent water - without adequate are the limiting factor. This is true, too, of limitlng
molsturetherecanbeno life. Water affectsland factors on mountains where lower altltudinal limits
anlmalsandplants in many ways. Airhumldity I S are set by heat or water or both, and upper altitudinal
Important In controlling loss of water through the limits are set by a lack of heat.
skm,lungs,and leaves. All
animals need some So Important are preclpitation and temperate that
form of water in thelr food or as drmk to run their several researchers use them to characterizebio-
excretorysystems.VascularplantshaveanInternal climates. Bioclimates are the aspects of climate that
plumbing system- parallel tubes ofdead tlssuecalled seem mostsignificant to livlngthlngs.Themost
xylem - that transfcrs water from root tlps to leaves. widely used bioclimatlc classification IS the 'climate
The entlre system is full of water under stress (capil- diagram'devlsed by HelnrlchWalter.This I S the

lary pressure). If thewater stress should fall too low, system of summarizmg ecophyslologlcal conditions
disaster may ensue - germlnation may Fail, seedlings that makes David Bellamy 'feel like a plant' (Rellamy
may not becomeestddishcd, and, should the fall occur 1976: 141)! Climate diagrams portray climate as a
durlng flowering, seed y~eldsmay be severely cut. An whole, Including the seasonal round of preclpitation
overlong drop In water stress kills plants, as anybody and temperature (Figure 2.6). They show at a glance
who has trledtogrowbeddingplantsdurlng a the annual patternof rainfall; the wet and dry seasons
drought and hose-pipe ban will know. characterlstlc of anarea, a s well as thelrIntensity,
since the evaporatlonrate 1s directly
related to
temperature; the occurrence or non-occurrence of a
Bioclimates
cold season, and the months in whlch early and late
O n land, precipitatron supplies water to ecosystems. frost have been recorded. Additionally, they provide
Plantscannot use allthepreclpitationthat falls. A informationonsuch factors as mean annualtem-
substantml portion of the precipltatlon evaporates and perature, mean annual preclpitation, the mean daily
returns to the atmosphere. For this reason, available mlnlmumtemperatureduringthecoldestmonth,
moisture (roughly the precipitationless the evapora- theabsoluteminimum recorded temperature,the
tion) is a better guide than precptatlonto the usable altitude of thestatlon,andthenumberof years
water in a terrestrial ecosystem. T h l s p o ~ nist readily of record.
understood wlth an example. A mean annual ramfall
of 400 mm might support a forest In Canada, where
W e t environments
evaporatlon is low, but mlght supporta dry savannah
in Tanzania, where evaporatlonIS high. Plants are very sensltive to water levels. Hydrophytes
Availablemoisturelargelydetermmes soilwater arewaterplantsandrootinstandingwater.Helo-
levels, which in turn greatly Influence plant growth. phytes are marsh plants. Mesophytes are plants that
For a plant to use energy for growth, water must be live innormally
molst but notwetconditlons.
available. Without water, the energy will merely Xerophytes are plants that live In dry conditions.
24 LIFE AND THE ENVIRONMENT

Wetlands support hydrophytes and helophytes.

O S M A N I Y E (120 m) 18.00~ 765
Thecommonwatercrowfoot (Rarrl/r/cnlu.c uqmf'tiis)
and the bog pondweed ( P o t m q e t o n pdyp/ifi,liI/.r) are
hydrophytes;thegreaterblrd's-foot trefoil (Lotm
/liigmu.ru.r) I S a helophyte. These plants manage to sur-
V I V ~by developlngasystemofalr spaces In their
roots, stems, o r leaves. The air spaces provide buoy-
ancyandimprove ~nternal ventilatlon. Mesophytes
vary greatly in their ability to tolerateflooding. In
thesouthernUnitedStates,bottomland hardwood
forestsoccupy swampsand rlver floodplains. They
contaln a setoftree specres that can survive In a
flooded habitat.Thewatertupelo (Ny.rstz q m f / u ) ,
which is found In bottomland forest in thesouth-
easternUnltedStates, is well adapted to suchwet
condicmns.
Flooding o r highsoil-moisture levels may cause
seasonal changes In mammal distributlons. The mole

ODESSA (70 m) 9.9"C392

[ l l - 251 F i g w e 2.6 Examples and explanation of climate diagram.
The letters denote the following. II Weather scaclon.
30 60 h Alt~tude (m above mean sea-level). L- Number of years o f
observatmn. Where there are two figures, the firsr refers to
temperature measurements and the secondtheprecipita-
20 40 rlon measurements. c/ Mean mnual temperature("C).
e Mean annual preclpcacion (mm).fMean tlaily maxlmum
10 20 temperaturedurlng the coldest month ("C). g A b s o l ~ ~ t e
minimum (lowest recorded) temperature ("C). 6 Curve of
mean monthlytemperature ( I scale graduarmn = IOOC).
-5.6 0
I Curve of mean monthly prec1p1tatwn ( 1 scale graduation
-25.0 = 2 0 mm). n / Relaclvely arld perlod or dry season (dotted).
11 Kelatwely humd perlod o r werseason(verrlcal bars).
HOHENHEIM (402 m) 8.4% 685 o Meanmonthly ralnfiall above 1 0 0 m m wlrh the scale
150 - 401 reduced by a factor o f 0 . 1 (the black area In Osmanlye).
p Curve for prec~pitationon a smaller scale ( 1 scale gradu-
40 80 atton = 3 0 mm). Above I C , horuontal broken lines mdicare
the relatively dry period or dry season (shown for Odessa).
30 60 q Monthswith a mean daily minimumtemperature
below 0°C (black boxes below zero line). r Months with an
absolute mlnlmumtemperature below 0°C (diagonal
20 40 lines). 5 Averageduration of period withdailymean
temperature above 0°C: (shown a s the number of days In
10 20 standard type); alternatlvely, the average dtmrion o f the
frost-free penod (shown :IS the number of days In Italic
type, a s for Honenhelm). M a n daily maxlmum tempera-
-3.5 0
-25.0 . turedurlng the warmestmonth ( h ) , absolute maximum
(hlghest recorded) temperature (i), :Ind meand;lily tem-
perature tluctuatlon ( j )are glven only for troplcal stations
with a diurnal climate, and are not shown In the eXampkS.
S o ~ o w :After Walter and I x r h ( 1960-7)
 LIFE AND THE ENVIRONMENT 25

rats (Cryptmys hottentotus) in Zimbabwe focus their Dry environments

activity around the bases of termite mounds during
Plants are very sensitive to drought and aridityposes
the rainy seasons as they rlse a metre or so above the
a problem of survival. Nonetheless, specles of algae
surroundinggrasslandandproducerelativelydry
growintheexceedinglydryGobidesert.Higher
islandsina sea ofwaterloggedterrain(Genelly
plants survlve in and conditionsby xerophytic adap-
1965).
tations - drylands support xerophytes. One means
Many organisms are fully adapted to watery envi-
of survival is slmply to avoid the drought as seeds
ronments and always have been - the colonnation of
(pluviotherophytes) or as below-ground storage organs
dry land is a geologlcal recent event. Some vertebrates
(bulbs, tubers, or rhizomes). Other xerophytic adap-
have returned to an aquatic exlstence. Those return-
tatlons enable plants to retain enough water to keep
ing to the water Include crocodiles, turtles, extinct
thelr protoplasts wet, so avoiding desiccation. Water
pleslosaurs and Ichthyosaurs, seals, and whales. Some,
is retalned by several mechanlsms. A very effective
including the otter,have adopted a semi-aquatlcway
mechanismiswaterstorage.Succulentsareplants
of life.
that store waterIn leaves, stems, or roots. The saguaro

Box 2.2
REPTILES IN DESERTS
Lizardsare abundantindesertsinthedaytime
whereas mammals are not. Thereason for this is not
a reduction of evaporative water loss through the
skin.Cutaneouswater loss is aboutthesamein
mammals and reptiles. However, reptiles from dry
habitats do have a lower skin permeability. There-
fore, they lose less water through the skin than do
reptilesfrommoistenvironments. The tropical,
tree-living green iguana (Iguana iguana) loses about
4.8 mglcm'lday through the skin and 3.4 mglcm2/
day through respiration; the desert-dwelling chuck-
walla (Sauromalrrs ohms), which is active in daytime,
loses about 1.3 mg/cm*/day through the skin and
1.1 mg/cm2/day through respiration. The difference
betweenmammalsandreptiles lies inthreerep-
tilian characteristics that predispose them for water
conservation in arid environments - low metabolic
rates; nitrogenous waste excretion as uric acid and
its salts; and, in many taxa, the presence of nasal salt
glands (an alternative pathway of salt excretion to
thekidneys). Low metabolicratesmean less fre-
quent breathing, which means thatless water is lost
from the lungs. Uric acid is only slightly soluble in
water and precipitates in urine to form a whitish,
semi-solid mass. Water is left behind and may be
reabsorbed into the blood andused to produce more
urine. This recycling of water is useful to reptiles
because their kidneys are unable to make urine with
a higher osmotic pressure than that of their blood
plasma. The potassium and sodium ions that do not
precipitate are reabsorbed in the bladder. This costs
energy, so why do it? The answer lies in a third
water-conservingmechanisminreptiles - extra-
renal salt excretion. In atleast three reptilian groups
- lizards,snakes,andturtles - therearesome
species that have saltglands.Theseglandsmake
possible the selective transport of ions out of the
body. They are most common in lizards, where they
have been found in five families. In these families, a
lateral nasal gland excretes salt. The secretions of
the glands are emptied into the nasal passages and
are expelled by sneezing and shaking of the head.
Salt glands are very efficient at excreting. The total
osmotic pressure of salt glands may be more than
six times that of urine produced by the kidney. This
explains the paradox of salt uptake from urine in
bladders. As ionsareactivelyreabsorbed, water
follows passively and the animalrecovers both water
andions from the urine. The ionscanthen be
excreted through the salt gland at much higher con-
centrations. There is thus a proportional reduction
in the amountof water needed to disposeof the salt.
26 LIFE AND THE ENVIRONMENT

Box 2.3
MAMMALS IN DESERTS from
air-dried
seeds.
Part of this
water
comes
from
the seeds and part comes from the oxidationof f d
- . . _ . . . .. I 1 I. \ I . I *
(metaDol1c water). Interestingly, tne water concent
Kodents are the dominant small mammals In arid
environments. Population densities may be higher of somedesertplants varies withtherelative
I:
t
t
ana .rive
. ..
~" tn" "nurrnws.
~ ~ _ "mm
~ n u.e~
~~ n nlenr-rune
"-
~ .
- ~ "
acuvlw
~ ~~, ~ ~

night be thought to avoid the heat stress of the day, The leavesareforagefor the oryx (Oryx gazeflu).
)eat stress can also occur at night when deserts can By feeding at night, the oryx takesin5litres of
.. - . , ... . ... . \ . . . I..-,. ._ .- _. &L L .""",
wacer. on wnlcn IC S U ~ V I V ~cnrouen
~ ~

S sevrral wacrr-

n the cool nasal passages. Overall,this process saves Hoarding food inaburrowprovidesnotonlya
water and energy. Indeed, the energetic savings are hedge against food shortages, but also an enhanced
10 great that it is unlikely that a homeotherm could source of water. In Texas,burrows of the plains
urvive without this system - ethmoturbinal bones pocket
gopher (Geomys hrrrsurius) have relative
n the fossil Cynognathus is persuasive evidence that humidities of 86-95 per cent (Kennedy 1964). In
nammal-like reptiles
(therapsids)
were
homeo- sealed burrows, the humidities can be up to 95 p e r
:berms. However, this countercurrent exchange of centwhilethe soil of theburrow floor contains
leatandmoisture is notanadaptationtodesert only 1 per cent water. Nonetheless, although high
.ife; it is an inevitable outcome of the anatomy and temperaturesand low humiditiesareavoidedin
?hysiology of the nasal passages. burrows, other stresses do occur - carbon dioxide
Water is also lost in faeces and urine. Rodents concentrations may be between ten and sixty times
3enerally can produce fairly dry faeces and concen- greater than in the normal air.
trated urine. Kangaroo rats, sand rats, and jerboas
:an produce urine concentrations double to quad-
ruple theurineconcentrationinhumans.The
spinifex hoppingmouse or dargawarra (Notomys
&xis), which lives throughout most of the central
snd western Australian arid zone, is a hot contender
For 'world champion urine concentrator'; its urine
concentration is six times higher than in humans
(Plate2.1). Low evaporativewater loss through
nocturnal habits, concentrated urine, and fairly dry
Plate 2.1 Spinifexhoppingmouse (N0)Omys
faeces mean that many desert rodents are indepen- -
alexis) world champion urine concentrator
dent of water - they can get all the water theyneed Photograph by Pat Morris
I-
 LIFE AND THE ENVIRONMENT 27

cactus (Curnegiea gzgunteu) andthe barrel

cactus
(Fwocactus uislizeni) areexamples. The barrelcactus
stores so much water thatit has been used as an emer-
gencywater supply by Indiansandotherdesert
inhabitants. Many succulents have a crassulacean
acid metabolism (CAM) pathway for carbon dioxide
assimilation.Thiskind of photosynthesis Involves
carbon dioxide being taken in at nlght wlth stomata
wideopen,andthenbeing used duringthe day
with stomata closed to protect against transpiration
losses. Other xerophytes (sometlmes regarded as
true xerophytes) do not store water, but have evolved
very effective ways of reduclng water loss - leaves
withthickcuticles,sunkenandsmallerstomata,
leaves shed duringdry periods, improvedwater
uptake through wide spreading or deeply penetrating
root systems,andimproved water conductlon.The
big sagebrush (Artemsiu tridentotu) is an example of a
true xerophyte. The creosote bush ( L r r e u divuricutu)
simply endures a drought by ceasing to grow when
water I S not available.
Plants vary enormously in theirabilityto with-
stand water shortages. Young plants suffer the worst.
Drought resistance is measured by the specific sur-
vival time(thetime between thepointwhenthe
roots can no longer take u p water and the onset of
dessicative injury).The specific survival time is
1,000 hours for prickly-pearcactus, 50 hours for
Scots pine, 16 hours for oak, 2 hours for beech, and
1 hour for forget-me-not (Larcher 1975: 172).
Desert-dwelling animals face a problem of water
shortage, as well as high daytime temperatures. They
have overcome these problems in several remarkable
ways (Boxes 2.2 and 2.3).
summer ranges andmovmgto lower elevations.
South-faclng slopes and windswept ridges, where snow
is shallower or on occasions absent, are preferred at
these times. In area of relatively level terrain, deer
and moose respond to deep snow by restrictmg their
actlvitiestoasmall area called 'yards' where they
establish trails through the snow. Prolonged winters
anddeep snow takea severe tollondeer andelk
populations.
For small mammals, snow is a blessing. It forms
an insulatingblanket,asort of crystallineduvet,
under which is a ground-surface mlcro-environment
where activity, including breeding In somespecles,
continuesthroughoutthewmter. To thesesmall
mammals, whlch Include
shrews (Sorex), pocket
gophers (Thonlort/.ys), voles (hficrotru, C/rthtronotny.r,
Phenucotnys), and lemm~ngs( k r t / t n u J , Dicrostonyr), the
most stressful tlmes are autumn, when intense cold
descends but snowfall has not yet moderated temper-
atures at the ground surface, and in the spring, when
rapid melting of adeep snowpackoftenresults in
local flooding. Another advantage of a deep snowpack
IS thatgreenvegetatlon may be available benrath
it, and several species make tunnels to gain access to
food.
Even in summer, snow may be important to some
mammals. Alpine or northern snowfields commonly
last through much of the summer on north-facing
slopes and provide acool microclimate unfavoured by
insects. Caribou (Rangijir turandus) and bighorn sheep
(Ovis spp.) sometimes congregate at these places to
seek relief from pesky warble flies.
The big picture: climatic zones

Snow Terrestrial ecozones

This is a significant ecological factor in polar and Onland,characteristicanimalandplantcommu-
some boreal envlronrnents. A snow cover that per- nlties areassociated with nine basic climatic types,
sists through the winter is a severe hardshlp to large varlously called zonobiomes (Walter 1985), ecozones
mammals. T o most North American artiodactyls, (Schultz 1995), and ecoregions (R. G. Bailey 1095,
including deer, elk, bighorn sheep, and moose, even 1996) (Figure 2.7):
moderate snow imposes a burden by covering some
food and making it difficult to find. In mountainous 1 Polar and subpolar zone. This zone includes the
areas,deer andelk avoid deep snow by abandoningArcticandAntarctic regions. It IS associated with
28 LIFE AND THE ENVIRONMENT
tundra vegetation. The Arctic tundra regionshave
low ramfall evenlydistributedthroughoutthe
year. Summers are short, wet, and cool. Winters
arelongandcold.Antarcnca is an icy desert,
although summer warming around the fringes is
causlng i t to bloom.
2 Boreal zone. This is thecold-temperatebelt
that supports coniferous forest(taiga). It usually
has cool,wetsummersand very coldwlnters
lasting at least six months. It is found only in the
NorthernHemispherewhere I t forms a broad
swath around the pole - it is a circumpolar zone.
3 Humid mid-latitude zone. Thls zone is the tem-
perate or nemoralzone.Incontinentalinteriors
I t has a short, cold winter and a warm, or even
hot, summer. Oceanic reglons, such as the Brltish
Isles, have warmerwintersandcooler,wetter
summers.This zone supports broad-leaved
deciduous forests.
4 Arid mid-latitude zone. This I S the cold-
temperate(conttnental)belt.Thedifference be-
tween summer and winter temperaturesis marked
and rainfall is low. Regions with a dry summer
butonly a slightdroughtsupporttemperate
grasslands. Reglons with a clearly defined drought
perlod and a short wet season support cold desert
and semi-desert vegetatlon.
5 Tropicalandsubtropicalarid zone. Thls I S a
hot desert climate that supports thorn and scrub
savannahs and hot deserts and semi-deserts.
6 Mediterraneansubtropical zone. Thls I S a belt
lylngbetweenroughly 35" and 45" latitude In
both hemispheres wlth winter rains and summer
drought. It supportssclerophyllous (thick-leaved),
woody vegetatlon adapted to drought and sensi-
twe to prolonged frost.
7 Seasonaltropical zone. Thls zoneextendsfrom
roughly 25" to 30" North and South. There IS a
marked seasonal temperaturedifference.Heavy
rain in the warmer summer period alternates with
extreme drought in the cooler winter perlod. The
annual rainfall andthedroughtperlod Increase
with distance from the equator. The vegetatlon is
tropical grassland or savannah.
8 Humid subtropical zone. Thls zone has almost
11o coldwinter season, and short wet summers. It
ISthe warm temperate climate In Walter's zono-
blome classlfication. Vegetation is subtroplcd
broad-leaved evergreen forest.
9 Humidtropical zone. This torrld zone has raln
all year and supports evergreen tropical r a n for-
est. The climate is s a d t o be diurnal because it
varies more by day and night than I t does through
the seasons.
Marine ecozones
The marinebiosphere alsoconsists of 'climatic'
zones, whlch are also called ecozones. The maln sur-
face-water marine ecozones arethe polar zone,the
temperate zone, and the troplcal zone (R. G. Bailey
1996: 161):
1 Polar zone. Ice covers thepolar seasIn winter.
Polar sras are greenish, cold,and have a low
salinity.
2 Temperace zone. Temperate seas are very mlxed
III character. They Include repons of high salinlty
111 the subtroplcs.
3 Tropical zone. Tropical seas aregenerallyblue,
warm, and have a high salinlty.
Biomes
Each ecozone supports severalcharacteristiccom-
munitlesofanlmalsandplantsknown as biomes
(Clements and Shelford 1939). The deciduous forest
biome in temperate western Europe is an example. It
consists largely of woodland wlth areas of heath and
moorland. A plantcommunityatthebiome scale
- all the plants associated with the deciduous wood-
land biome, for example - is a plant formation. An
equivalent animal community has no specla1 name; it
is simply an anlmal community. Smaller communltm
wlthln biomes are usuallybased on plant distribution.
They arecalled plantassociations. InEngland,
associations wlthin the deciduousforest blome Include
beech forest,lowlandoakforest, andash forest.
Between h o m e s aretransitlonalbeltswhere the
climate changes from one type to the next. These are
called ecotones.
'p
5
I
30 L I F E AND T H E ENVIRONMENT

Zonobiomes tered beech (FaguJ Jyhatrca) trees at lower elevations.

Mountain pines (Pinus montana) with scattered Swiss
All the biomes around the world found in a stone pines (Pinus cembra)grow near the tree line. The
particular ecozone constitute a zonobiome. A plant subalpinebelt lies between about 3,000 mand
communlty at the same large scale is a formation- 3,500 m.Itcontamsdiminutive forests of tiny
type or zonal plant formation. The broad-leaved willow (Salix spp.) trees, only a few centimetres tall
temperate forests of western Europe, North America, when mature, wlthin an alpine grassland. The alpine
eastern Asla, southern Chile, south-east Australia and belt, which extendsup to about 4,000 m, is a
Tasmanla,andmost of New Zealand comprisethe meadow of patchy grass and a profusion of alpine
humidtemperate zonobiome.Between the zono- flowers - popples,gentians, saxifrages, and many
biomes are transitional belts
where the
climate more.Themountaintops above about 4,000 m
changes from one type to the next. These are called liewlchln thenivalzoneand are covered with
zonoecotones. permanent snow and ~ c e .
Freshwater communlties (lakes, rivers, marshes,
andswamps) are part of continental zonobiomes.
They may be subdivided In various ways. Lakes, for SOILS,SLOPES, AND DISTURBING
Instance, may be wellmlxed (polymictic o r oligo- A G E N C I E S : P E D O G E O M O R P H I C
mictic) o r permanently layered (meromictic). They F A C T O R S A N D D I S T U R B A N C E
may be wantlng in nutrientsandbiota(oligo-
trophic) or rich in nutrients and algae (eutrophic).
A
thermocline(wherethe
temperature profile Soil and substrate
changes most rapidly) separates a surface-water layer Soil and substrate influence animals and plants, both
mixed by wind (epilimnon) from a more slugglsh, at the level of individual species and at the level of
deep-water layer (hypolimnon). And, as depositional communities.
environments, lakes are divided into a littoral (near-
shore) zone, and a profundal (basinal) zone.
Marine ecozones, and the deep-water regions, con- Microbes and substrate
sist of biomes (equivalent to terrestrial zonobiomes). Acid-loving microbes (acidophiles) prosper in
The chief marine biomes are the intertidal (estuar- environmentswithapH below 5 . Sulfolobus acido-
ine, littoralmarine, algal bed, coral reef) biome, caldarius, as well as liking it hot, also likes it acid.
the open sea (pelagic) biome,theupwelling zone Alkali-loving microbes (alkaliphiles) prefer an envi-
biome, the benthic biome, and the hydrothermal ventronment with apH above 9. Natronobacterirm2 gregotyi
biome. lives in soda lakes.
Salt-loving microbes live intensely
in saline
environments.
They survive by producing large
Orobiomes
amounts of internal solutes that prevents rapid de-
Mountain areas possess theirown biomescalled hydration in a salty medium. An
example is
orobiomes. The basic environmental zones seen on Halobactevium salinarintn.
ascending a mountain are submontane (colline, low-
land),montane,subalpine,alpine,and nival. O n Plants and substrate
south-facing slopes in the Swiss Alps around Cortina,
the submontane belt lies below about 1,000 m. It Plants seemcapable of adapting to the harshest of
consists of oak forests and fields. The montane belt substrates.Saxicolousvegetationgrows on cliffs,
ranges fromabout 1,000 m to 3,000 m. The bulk rocks, and screes, some species preferring rock crevices
of it is Norway spruce (Pic-eaabies) forest, with scat- (chasmophytes), othersfavouring small ledges where

detritus and humushave collected (chomophytes). In
the Peak District of Derbyshire,England,maiden-
hair spleenwort (AspIeniunI tric%lomanes)I S a common
chasmophyte and thewallflower (Chriranthus chnri) is
common
a and colourful chomophyte(Anderson
and Shlmwell 1981: 142). Perhaps the most extreme
adaptationto a harsh envlronment is seen in the
mesquite trees (Pro.ropis tamarqo and ProsopcJ alhu)
that grow In the Pampa del Tamagural, aclosed basm,
or salar, in the rainless region of the Atacama Desert,
Chile. These plants manage to surviveon concrete-
like carbonate surfaces (Ehleringer et a / . 1992). Thelr
leaves absclse (are shed) and accumulate to depths of
45 cm. Because there I S virtually no surface water, the
leaves do notdecompose and nitrogen I S not Incor-
porated back Into the soil for recycling by plants. The
thick, crystalline pan of carbonate salts prevents roots
from growing Into the litter. To survlve, treesthe have
roots that tix nitrogen i n m o m subsurface layers, and
extract moisture and nutrients from groundwater at
depths of6-8 mor more through a taproot and amesh
of fine roots lymg between 50 and 200 cm below the
salt crust. A unique feature of thls ecosystem is the
lack of nitrogen cycling.
Calcicoles (or calciphiles) are plants that favour
such calcium-rich rocks as chalk andlimestone.
Calcicolous species often grow onlyonsoilformed
inchalk or limestone.Anexample from England,
Wales, and Scotland is the meadow oat-grass (Helicto-
trichon pratense), the distribution of which picks out
the areas of chalk and limestone and the calcium-rich
schists of theScottishHighlands.Other examples
are traveller’s joy (Clematis vitalba), the spindle tree
(Euonymus europaeu), andthe
common rock-rose
(Helianthemmnummularium). Calcifuges (or calci-
phobes) avoid calcium-rich soils, preferring instead
acidic soils developed on rocks deficient in calcium.
An example is the wavy hair-grass (Desrhampsia
flex/losa). However, many calcifuges
are
seldom
entirely restricted to exposures of acidic rocks. In the
limestone Pennine dales, the wavy hair-grass can be
found growing alongside meadow oat-grass. Neutro-
philes are acidity ‘mlddle-of-the-roaders’. They
tend to grow in the range p H 5-7. In the Pennine
dales, strongly growing, hlghly competitive grasses
LIFE AND T H E ENVIRONMENT 31
thatmake heavy demandsonwaterandnutrient
stores are the most common neutrophiles.
Animals and substrate
Some animals are affected by soil and substrate. For
instance, the type and texture of soil or substrate is
critlcal to twokmds of mammals: those that seek
diurnal refuge In burrows, and those that have modes
of locomotlonsuitedto relatively rough surfaces.
Burrowing species, which tend to be small, may be
confined to a particularkind of soil. For instance,
many desert rodents displaymarkedpreferences for
certaln substrates. In most deserts, no slngle species
of rodent I S found on all substrates; and some species
occupyonlyone substrate. Fourspecles of pocket
mice (Peropathu.c) live in Nevada,UnltedStates
(Hall 1946). Their preferences for soil types are
largely complementary: one lives on h r l y firm soils
of slightlysloping valley margins;the second is
restricted to slopes where stonesand cobblesare
scatteredandpartlyembedded in theground;the
thlrd is associated wlththe fine, silty soil of the
bottomland;andthefourth,asubstrategeneralist,
can survive on a variety of soil types.
Saxicolous species grow in,or live among, rocks.
Somewoodrats (Neotoma) buildtheir homesexclu-
sively in cliffs or steep rocky outcrops.The dwarf
shrew (Sorex nanrrs) seems confined to rocky areas in
alpine and subalpine environments. Even some salt-
atorial species are adaptedto life on rocks. The
Australian rock wallabies (Petrogale and Petrodorcas)
leap adroitly among rocks (Colour plate 1). They are
aided in this by traction-increasing granular patterns
on the soles of their hind feet. Rocky Mountain pikas
(Ochotona prrnreps) in the southern Rocky Mountalns,
UnitedStates, normallylive ontalus or extensive
piles of gravel (Hafner1994).Thoseliving near
Bodie, a ghost townin the Sierra Nevada, utilize tail-
ings of abandonedgoldmines(A. T. Smith 1974,
1980). The yellow-bellied marmot (Marmota&viven-
tris) is another saxlcolous species, andcommonly
occurs with the Rocky Mountain pika. The entirelife
style of Afrlcan rock hyraxes (Hetwohyrax, Procavra) IS
built around their occupancy of rock piles and cliffs.
32 LIFE AND THE ENVIRONMENT

Most of thelr food consists of plants growing among, Altitude

or very close by, rocks. Thelr social system is bonded
Altitude exertsastronginfluence onanlmalsand
by thescent of urine and faeces on the rocks. The
plants. The plant communities girdling the Earth as
rocks provldeusefulvantagepoints to keep an eye
broad zonal beltsare paralleled In theplant com-
out for predators, hlding places, and an economical
munities encircling mountains - orobiomes (17. 30).
means of conservmg energy.
Individualanimalsandplant speciesoftenoccur
withln a partlcular elevational band, largely owlng to
Pedobiomes climaticlimlts of tolerance.However,altitudinal
rangesare Influenced by ahostofenvironmental
Withln zonobiomes,thereareareas of intrazonal
factors, and not lust climatic ones. For instance, tree-
andazonalsoilsthat, in some cases, supporta
linesarenot always purely the result of climatic
distinctlvevegetation.These non-zonal vegetation
constraints on tree growth, but involve pedological
communlties are pedobiomes (Walter and Breckle
and blotic lnteractlons as well.
1985). Several different
pedobiomes are
distln-
guishedonthe basis of soiltype:lithobiomeson
stony soil, psammoblomes on sandy soil, haloblomes Aspect
on salty soil, helobiomes in marshes, hydrobiomes on Aspect (compass direction) strongly affects the
waterlogged soil, peinoblomes on nutrrent-poor soils, climate just above the ground and within the upper
andamphiblomeson soils thatare flooded only soillayers. For thls reason,virtuallyalllandscapes
part of the time ( e . 6 , river banksandmangroves). display significant differences in soil and vegetation
Pedobiomes commonly form a mosaic of small areas onadjacentnorth-facing(distal)andsouth-facing
and are found in all zonoblomes. There are Instances (proximal) slopes, and on the windward and leeward
wherepedoblomes areextensive: theSuddmarshes sides of hills and
mountains. In
the
Northern
on the Whlte Nile,whlch cover 150,000 km'; fluvlo- Hemisphere, south-faclng slopes tend to be warmer,
glacial sandy plalns; andthe nutrient poorsoils of the and so moreprone to drought,than north-facing
Campos Cerrados in Brazil. slopes. The difference may be greater than imagined.
A striklng example of a lithobiome is found on In aDerbyshiredale,England,thesummermean
serpentine. The rock serpentine and its relatlves, the temperature was 3°C higher on a south-facing slope
serpentinites, are deficient in :duminium. This leads than on a north-facing slope (Rorison et ai. 19861, a
to slow rates of clay formatlon, whlch explains the difference equivalent to a latltudinal shiftof hundreds
characterlstic fratures ofsoils formed on serpentlnltes: of kilometres! These differences affect plant growth.
they are high erodible, sllallow, and stock few nutrl- The LBgern is an east-west trending mountain range
ents.Thesepeculiarfeatureshaveaneye-catching near Baden in the SwissJura.Cool north-faclng slopes
Influence onvegetation(Brooks1987;Baker et u/. areseparatedfromwarmsouth-facingslopes by a
1992).Outcrops of serpentine support small Islands half-metre-wldemountalnrldge.Thesouth-facing
of brush and bare ground In a sea of forest and grass- slopesupportswarmth-lovlngvegetation - downy
land. These islands are populated by native floras with oak (Qlterczrs p t t h c ~ ~ ~ 'woodland
~.c) with pale-flowered
many endemic spec~es (Whlttaker 1954). orchid (Orcha pu//ens). wonder violet (Violu vtiruhiiis),
and bastard balm (/2lriittzsrtte/i.csoph~iitrrt~).
The north-
fiaclng slope supports a beech (Fagz/.r .ryizwrrtzcu) forest
The ups and downs of living:
wlthsuchsubalpineplantsasalplne penny-cress
topography
(Th/u.rpi uipe.rtrr) andgreenspleenwort (As,bietti/trr1
Many topographic factors Influence
ecosystems. tvrzde) (Stout~esdijkandBarkman1992:78).The
Themost rnfluentialfactorsare altitude,aspect, change over the narrow mountain r d g e I S equivalent
inclination, and insularity. botanically to about 1,000 km of latitude.
 L I F E AND THE ENVIRONMENT 33

The geographicaldistributions of someanimal cotton-grass (Eriopborrrtt/ anq/.rtlfo/iurrt) becoming

spec~esare influenced by aspect. O n the south-faclng more important towards the bottom end. Peaty gley
slopes of the Kullaberg
Penlnsula, south-west soils and peat soils form in the wet lower slopes and
Sweden, several arthropod species are found far to the supportpurplemoor-grass,commoncotton-grass,
north of their maln rangc. The spec~es include the andbog mosses (Sphagnutn spp.),withtussocks of
moth Idura cfillrtaria, the beetle Danal.ea pa/fipe.r, and hare’s-tail cotton-grass (Errophor/rm z~aglnatuw) in
the spder ‘I;herrdion c o n r p w n (Ryrholm 1988). Also bogs and rushes (J//tmr.r spp.) In acld pools of water.
In Europe, Alpine marmots ( h f a w o t amartt/ota) prefer Topographic factorsnormallyworkintandem
to burrow on s o u t h - k i n g slopes, although summer withother factors. This is seen at smallandlarge
temperatures on theseslopes are high enough to limit scales. O n the small scale, aspect and topographical
theirfeedingtimeabovegroundandnorth-facing posltlon are good predictors of funnel ant (Aphaeno-
slopes would seem a wlser cholce.They may dig p t lungzeps) ~ nests In a20-haeucalypt-forestsite
burrows in south-Faclng slopes because conditlons for at WogWog, south-eastern NewSouth
Wales
hibernating are better (Tiirk and Arnold 1988). (Nicholls and McKenzle 1994). O n a large scale, ele-
Differences between the climate of windward and vatlon and
altltudinal
rangepartly
explainthe
leeward slopesmayalso beconsequential. Large distribution of sixty-onewillow (Salix) specles in
mountaln ranges cast a rain shadow In thelr lee that Europe, though latltude and July mean temperature
is sufficlent to alterthevegetation.Innorth-west have somewhat more explanatory power (Myklestad
NorthAmerica,thewet coastal mountalnranges and Birks 1993).
support luxurlant
Sitka
spruce (Pil.eu s ~ t t b e t ~ ~ l s ) ,
Douglas fir (PseIrdotsIqa merrzte.rii), and hemlock
Insularity
(Tsuga spp.) forests. Furtherinland, In the rain
shadow of the Cascade Mountalns, the valley of the Insularity I S the relative Isolation of an ecosystem.
Columbia Rlver is a desert supportmg sagebrush and It has asignificant Influence onIndividualsand
tufts of drought-resistant bunch grasses. communitles.Trueislands have abnormallyhigh
rates of extinction, house many endemic and relict
species, favour theevolutlonofdwarf,giant,and
Inclination
flightlessforms,andareofteninhablted by good
Slope is the inclinatlon of the ground from the hor- dispersers. Habitat Islands - woods withln fields,
izontal. It affects vegetation through its influence on remnant patches of heathland, mountain tops, parks
soil moisture and on substrate stability. Slope com- In cltles,and so on - arebecomingincreasingly
monly varies in a regular sequence over undulating common as the land cover becomes more and more
terraln - from summit, down hillside, to valley bot- fragmented. The survlval of populationsin fragment-
tom - to form a geomorphic catena or toposequence. ing habitat islands is currently belng assessed using
The geomorphlc toposequenceis mirrored In soil and metapopulation models (Chapter4).
vegetation toposequences. Figure 2.8 shows a soil
andvegetatlontoposequence onHodnet Heath,
Disturbance
Shropshire,England.Humus-Ironpodzolsformon
drysummit slopesandsupporteitherdryheather Disturbance is any event that disrupts an ecosystem.
(Call1mu zudgar/J) heathor silverblrch (Betnla pen- Disturbingagencies may be physlcal orblological.
d d a ) wood wlth bracken (Ptrridim aqlrilinilrtt/) and Grazing, for example, disturbs vegetation communi-
wavy hair-grass (DeJchatt/pm fixnosa). Gley podzok ties, although I t origlnates within the same ecosystem
formonthewettermld-slopesandsupportdamp astheplantsthat it disturbs. It is usefd to thlnk
cross-leaved heath ( E r m trfralix) and heather, with ofdisturbance as a contlnuum between purely external
purplemoor-grass (c\.lo/inra mrrrlea) andcommon agenclesand
purely
internal
agencies. Classical
34 LIFE AND THE ENVIRONMENT

I I
Dry heather heath or
birch wood with bracken
and wavy hair-grass
I I
-
Silver birch (Betula pendula)
I I 1
I Damp heather-cross-leaved heath I Purple moor-grass, common
I heath with purple moor-grass and I cotton-grass and mosswith
I common cotton-grass increasing I hare's-tail cotton-grass (bog)
I towards the wetter end I and rushes (acid pools)
I
""I

Heather (Calluna vulgarrs)

.-M "

8 Cross-leaved heath (Erica tetralix)

n
v1
I

'c
0 """"I

W
M
C ""I-

.............

.........uii.. "
..................
""~~,;g.:~.~.:.~.:..,~.:::,::f::.:
.. .-...-..+....
. . . . .......................
. . . .
.................. . .
L
10 m I

I I I
-VI
.- I
II
Humus-iron podzols I Gley podzols I
I
Peaty
gley
soils Peat
soils
v)
I I

P i p r e 2.8 Soil and vegetation toposequences on Hodnet Henrh, Shropshlre, England. The letters L, F, H. etc. are soil
horizons.
Sorow: After Burnham and Mackney (1968)

'disturbance' lies at the externalagency end of the disturbmg agenciesonecosystems can be dramatic.
continuum. I t IS the outcome of p h y s d processes Pathogens, for example, are forceful disrupters of
actingata particular time,creatingsharppatch ecosystems at small and medium scales, witness the
boundaries,
and increaslng resource availability. efficacy of Dutchelm disease in Englandandthe
Internal community cycles, such as the hummock- chestnutblight In the Appalachian regioneastern
hollow cycle In peatland, lie at the Internal end of Unlted States (p. 80).
the disturbance continuum and are driven by Internal Some disturbances act essentially randomly within
community dynamics. a landscape to produce disturbance patches. Strong
Ecosystems may be physically disturbed by strong windscommonly behave In thls way. Thepatches
winds(whichuproot trees), fire, flood, landslides, produced by randomdisturbance can be extenswe:
andlightnlng;and biotically disturbed by pests, the blomantles formed by burrowing animals are a
pathogens, and the activities of animals and plants, case in point (D. L. Johnson 1989, 1990), as are the
includingthehuman specles. The effects of these patches of eroded soil created by grizzly bears (Urws

arct0.r t5orrihili.r) excavatlngdens,digging for food,
andtrampling well-established trails(Butler1992;
see also Butler1995). Second, somedisturbances,
such as fire, pests, and pathogens, tend to start at a
pointwithina landscape andthen spread toother
parts.Inboth cases, thedisturbancesoperate In a
heterogeneousmanner because some sltes within
landscapeswill be more susceptibletodisturbance
agencies than others.
Wind
Tree-throw by strongwinds,andto lesser extent
other factors, may have a considerable impact on
communities.In manyforests, disturbance by up-
rooting I S the primary means by whlch species rich-
ness is maintained (Schaetzl et al. 1989a, b). A fallen
tree creates agap in the forest that seemsvital
to community and vegetation dynamlcs and succes-
slonal
pathways: I t provides
niches wlth
much
sunlight for pioneerspecies,encourages the release
of suppressed, shade-tolerant saplings, and aids the
recrultment of new inclividuals. Even tropical forests,
once thought Immune to physical disturbance, have
been shown to contaln seemingly haphazard patterns
of treeforms, age classes, and specles. The troplcal
rain forests of the Far East may be analysed as ‘gap
phases’ (Whitmore 1975). Gaps are openings made
In the forests by varlous disturbances, and the phases
are the stages of treegrowth In thegaps, from
seedlingtomaturityanddeath. Small gaps,about
0.04 ha, are opened up by the fall of Individual large
trees, wlth crowns 15-18 m in diameter. Llghtnlng
strikes open up gaps with an area of about 0.6 ha.
Local storms cause larger gaps (up to 80 ha), while
typhoonsand tornadoes destroy even larger areas.
These gaps are an Integral part of the tropical forest
system.
Fire
Fire is relatively common in many terrestrial
envlronments.It influences thestructureandfunc-
tlon of someplant species and many communities
(e.g. Whelan 1995). The effects of tire are beneficial
LIFE AND THE ENVIRONMENT 35
and detrmental. There are three main detrmental
effects. First, many organisms are lost from the com-
munity. Second, after a severe fire, the ground is left
vulnerable to soil eroslon. Third, minerals are lost In
smokeandthroughvolatilization,which releases
large quantities of nitrogen and sulphur. Beneficial
effects are that dead litter is burnttoash, which
boosts mineral recycling, andthatnitrogen-fixing
legumes often appear in the aftermath of a moderate
surface (asopposed tocrown) fire. In areas where
fires are frequent, many plants are tolerant of tire,
and some require a tire to prompt seed release and
germination. In the Swartboskloof catchment, near
Stellenbosch, Cape Province, South Africa, mountain
fynbos (Mediterranean-type)plants possess awide
range of regeneratlonstrategiesand fire-survival
mechanisms (van Wilgen et al. 1992). Most species
can sproutagam after a tire, and areresilient to
a range of fire regimes. Few species are reliant solely
on seeds for regeneratlon.Thosethat are obligate
reseeders regenerate from seeds stored In the soil, and
just a few, such as Protu neriifolia, maintain seed
stores In the canopy. Most of the fynbos plants flower
withm In a year of a tire. Somespecies, including
Wdtsonia horhonicu and Cyrtrmth~~rventrtt-osm, are
stimulated into flowering by tires, though the trlgger
appears to act indirectlythroughchanges in the
environment (such as altered soil temperature
regmes), and not directly through heat damage to
leaves or aplcal buds.
A D A P T I N G TO C I R C U M S T A N C E S :
N I C H E S A N D LIFE-FORMS
Ways of living
Organisms have evolved to survive In the varied
conditlons found at theEarth’s surface.They have
come to occupy nearly all habltats and to have filled
multifarious roles withln food chalns.
Ecological niche
An organism's ecological niche (or simply niche) I S
36 LIFE AND THE ENVIRONMENT
its ‘address’ and ‘profession’. Its address or home IS
the habitat in which I t lives, and is sometimes called
the habitat niche. Its professlon or occupation IS its
position ina food chain,and is sometlmes called
the functional niche. A skylark’s (Alarrda arvensis)
address I S open moorland (and, recently, arable farm-
land); its profession is insect-cum-seed-eater. A
merlin’s (Falro cohnbarttrs) address is open country,
especially moorland;its profession I S a bird-eater,
withskylarkand meadow piplt (Anthus pratenszs)
bemg its main prey. A grey squirrel’s (Sciurtrs caro-
(inensis) habitatnlche is a deciduouswoodland; Its
profession is anut-eater (smallherbivore). A grey
wolfs (Cants II@J) habitatnlche is cool temperate
coniferous forest, and Its profession is large-mammal-
eater.
A distinction IS drawn between the fundamental
niche and the realized niche. The fundamental (or
virtual)nichecircumscribes where anorganism
would live underoptimal physical conditionsand
with no competitors or predators. The realized (or
actual) niche IS always smaller, and defines the ‘real-
world’ niche occupled by an organism constrained by
biotic and abiotic limiting factors.
A niche reflects how an Individual, species, or pop-
ulation interacts wlth and exploits its envlronment.
lnvolves adaptation to
environmental conditions.
The competitive exclusion principle (Chapter 5)
precludes two species occupyingidentical niches.
However, a group of species, or guild, may exploit
the same class of envlronmental resources in a similar
way (R. B. Root 1967; Simberloff and Dayan 1991).
In an oak woodland,oneguild of birds forages
for arthropods from the foliage of oak trees; another
catchesinsects In theair;anothereats seeds. The
foliage-gleaning guild in a California oak woodland
includes members of four families: the plain tltmouse
(Prrrm inornntzrs, Paridae), the blue-gray gnatcatcher
(Po/ioptila caertrlea, Sylviidae), the warbling vireo and
Hutton’s vireo (Vireo gilvrtr and Vireo huttoui, Vireo-
nldae),andtheorange-crowned warbler (Vwmizwu
d a t a , Parulidae) (R. B. Root 1967).
Ecological equivalents
Although each niche is occupied by only one species,
different species may occupy thesame or slmilar
niches in different geographical regions. These species
are ecological equivalents or vicars.A grassland
ecosystem contains a niche for large herbivores living
in herds. Bison and the pronghorn antelope occupy
this nichein North America; antelopes, gazelles,
zebra, and eland In Africa; wild horses and asses in
Europe;thepampasdeerandguanaco in South
America; and kangaroos and wallabies In Australia.
As thisexample shows, quitedistinct species may
become ecological equivalents through hlstorical and
geographical accidents.
Manyblrd guilds have ecological equivalents on
different continents.Thenectar-eating(nectivore)
guild has representatwes in North Amerlca, South
America,and Africa. InChileand California the
representativesare the hummingbirds (Trochilidae)
andthe African representativesare thesunbirds
(Nectariniidae). One remarkable convergent feature
between hummingblrds and sunbirdsis the iridescent
plumage.
Plant species of very different stockgrowlng in
Itdifferent areas, when subjected to the same environ-
mental pressures, have evolved the same life-form to
fill the same ecologicalniche. The American cactus
and the South African euphorbia, both living in arid
regions, have adapted by evolvlng fleshy, succulent
stemsand by evolving spines Instead of leaves to
conserve precious moisture.
life-forms
Thestructureand physlology of plantsand,toa
lesser extent, anlmals are often adapted for life in a
particular habitat. These structural and physiological
adaptatlons are reflected in life-form and often
connected with particular ecozones.
Thelife-form of anorganism is its shape or
appearance, its structure, its hablts, and Its kind of
life hlstory. I t includes overall form (such as herb,
shrub, or tree in the case of plants) and the form of
individual features (such as leaves). Importantly, the

dominant types of plant In each ecozone tend to have
a life-form finely tuned for survlval under in that
climate.
Plant life-forms
A wldely used classification of plantlife-forms,
based on the position of the shoot-apices (the tips of
branches)where new buds appear, was designed by
ChristenRaunkiaer in 1903 (see Raunkiaer1934).
distingulshes
It five maln groups: therophytes,
cryptophytes, hemlcryptophytes, chamaephytes, and
phanerophytes (Box 2.4).
A biological spectrum IS the percentages of the
differentlife-forms In agiven reglon. The‘normal
spectrum’ IS a kind of reference polnt; it I S the per-
centages of differentlife-forms in the world flora.
Each ecozone possesses a characteristlcblologlcal
spectrumthat differs from the‘normalspectrum’.
Tropical forests containa wide spectrum of life-
forms, whereas in extreme climates, with either cold
or dry seasons, the spectrum 1s smaller (Figure 2.10).
As a rule of thumb, very predictable, stable climates,
such as humid tropical climates,support a wider
variety of plant life-forms thando regions wlth
inconstant climates, such as arid, Mediterranean, and
alpineclimates.Alpine regions, for instance, lack
trees, thedominant life-form belng dwarf shrubs
(chamaephytes).IntheGrampianMountains,Scot-
land, 27 per cent of the specles are chamaephytes, a
figure threetimesgreaterthanthepercentage of
chamaephytes in the world flora (Tansley 1939).
Some life-forms appear to be constrained by climatic
factors. Megaphanerophytes (where the regeneratmg
partsstand over 30 m from the ground) arefound
only where the mean annualtemperature of the
warmest month I S 10°C or more. Trees are confined
to places where the mean summertemperature
exceeds 10”C,bothaltitudinallyandlatitudinally.
Thlsuniform behaviour I S somewhatsurprising as
different taxa are Involved in differentcountries.
Intrlguingly, dwarf shrubs, whose life cycles are very
similarto those of trees, always extendtohigher
altitudes and latltudes than do trees (Grace 1987).
Indivldual parts of plants also display remarkable
L I F E A N D THE ENVIRONMENT 37
adaptations to life in different ecozones. This I S very
true of leaves. Inhumldtropical lowlands, forest
trees have evergreen leaves wlth no lobes. In regions
of Mediterranean climate, plants have small, sclero-
phyllous evergreen leaves. Inarid regions, stem
succulents without leaves, such as cactl, and plants
withentlre leaf margins (especially among ever-
greens) have evolved. In cold wet climates,plants
commonly possess notched or lobed leaf margms.
Animal life-forms
Animal life-forms, unlike those of plants, tend t o
matchtaxonomic categorles ratherthan ecozones.
Most mammals are adapted to basic habltats and may
be classlfied accordingly.They may be adapted for
life in water (aquaticor swimming mammals), under-
ground(fossorial or burrowingmammals), on the
ground(cursorial or running,andsaltatorial or
leapingmammals), In trees (arboreal or climbing
mammals), and in the air (aerial or flying mammals)
(Osburn etal. 1903). None ofthese habitats is strongly
related toclimate.That is not to say thatanlmal
species arenot adaptedtoclimate:there aremany
well-known cases of adaptation to marginal environ-
ments, including deserts (pp. 25-6) (see Cloudsley-
Thompson 197 5 b).
Autoecological accounts
Detailed habltat requirements of Individual species
require careful and intensive study. A ground-break-
ing study was the autoecological accountsprepared
for plantsaround Sheffield, England(Grime et al.
1988).About 3,000 km’ were studied in three sepa-
ratesurveys by theNaturalEnvironment Research
Council’s Unit of ComparativePlant Ecology (for-
merly theNature ConservancyGrassland Research
Unit).The regioncomprises tworoughly equal
portions: an ‘upland’ reglon, mainlyabove 200 m and
with mean annual precipitation more than 850 mm,
underlain by Carboniferous Limestone, Millstone
Grit, and Lower Coal Measures; and a drier, ‘lowland’
region overlying Magnesian Limestone, Bunter
Sandstone, and Keuper Marl.
38 LIFE AND THE ENVIRONMENT

Lox 2.4
'LAN1 LIFE FORMS subtypes - herbaceous
phanerophytes,
epiphytes,
and stem succulents. A herbaceous example is the
scaevola, Scaevola koenigii. Phanerophytes are divided
'hanerophytes into four size classes: megaphanerophytes (> 30 m),
. .

risible) are trees and large shrubs. They bear their (2-8 m) and nanophanerophytes (c 2 m).
)udsonshoots that projectintotheairandare
lestined to last many years. The buds ate exposed
Chamaephytes
o the extremes of climate. The primary shoots, and
n many cases the lateral shoots as well, are nega- Chamaephytes (from the Greekkhamai, meaning on
ively geotropic (they
stickup
into
the
air). theground)aresmallshrubs,creepingwoody
Veeping trees are an exception. Raunkiaer (1934) plants, and herbs. They bud from shoot-apices very
livided phanerophytes into twelve subtypes accord- close to the ground. The flowering shoots project
ngtotheirbudcovering(withbud-covering or freely into the air but live only during the favourable
without it), habit (deciduous or evergreen), and size season. The persistent shoots bearing buds lie along
mega, meso,micro,andnano);andthreeother the soil, rising no more than 20-30 cm above it.

I
Hemicrypto- ' Therophytes
Crypfophytes
Phanerophytes
Chamaephytes
bhvtes
,- I

Surviving buds or Surviving buds or Surviving buds or Surviving buds or shoot-apices

survive
shoot-apices borne shoot-apices very shoot-apices at buried in ground -tuberous seedsas
on geostrophic close to, but above, soil surface - herbs and bulbous herbs
shoots projeding ground - creeping growing rosettes
-
into the air woody plants and tussocks
shrubs and trees and herbs

Figure 2.9 Plant life-forms.

Sowce: AfterRaunkiaer (1934)
 L I F E AND THE ENVIRONMENT 39

Suffructicose chamaephytes have erect aerial shoots and bear foliage leaves) such as the vervain (Vwbena
that die back to the ground when the unfavourable oficinalis), partialrosetteplantssuch as the bugle
season starts. They include species of the Labiatae, (Ajuga reptans),and rosette plants such as the daisy
Caryophyllaceae, and Leguminosae. Passive chamae- (Bellisperennis).
phytes have procumbentpersistentshoots - they
are long, slender, comparatively flaccid, and heavy
Cryptophytes
and so lie alongthe ground. Examples are the greater
stitchwort (Stellaria hoIostea) and the
prostrate Cryptophytes are tuberous and bulbous herbs. The) I

speedwell (Veronica postratu). Active chamaephytes areeven more ‘hidden’thanhemicryptophytes -

have procumbent persistent shoots that lie along thetheir buds are completely buried beneath the soil
ground becausetheyaretransverselygeotropic in thus affording them extra protection from freezing
light (take up a horizontal position in response to and drying. They include geophytes (with rhizomes
gravity). Examples are the heath speedwell (Veronica bulbs, stem tubers, and root tuber varieties) such a!
ofiinulis), thecrowberry (Empetrumnigrurn), and the purple (Crmris
crocus m u s ) , helophytes or marst
the twinflower (Linnaea borealis).Cushion plants are plants such as the arrowhead (Sagittariasagittifolia)
transitionaltohemicryptophytes.They have very and hydrophytes or water plants such as the rootec
low shoots, very closely packed together. Examples shining pondweed (Potamogeton lrrcens) and the free,
are the hairy rock-cress(Arabis hirusa)and the house- swimming frogbit(Hydrocharis nzwsrrrr-ranae).
leek (Semptwivum tatwarn).
Therophytes
Hemicryptophytes
Therophytes (from the Greek tberos, meaning sum
r .

hidden) are herbs growing rosettesor tussocks. They favourable season and survive the adverse season a
budfromshoot-apiceslocatedinthesoilsurface. seeds. Examples are the cleavers (Gulirrm aparine)
They include protohemicryptophytes (from thebase the cornflower (Centaurea cyanuf), andthe wal
upwards, the aerial shootshave elongated internodes hawks-beard (Crepis tectorurn).

An example of the ‘autoecological accounts’ forthe vulnerabletograzing,cutting, or trampling.Its

bluebell (Hyacznthoider non-scrzpta) is shown in Figure
2.1 1. The bluebell is a polycarpic perennial, rosette-
formmggeophyte,withadeeplyburiedbulb.It
appears above ground In the spring, when I t exploits
the light phasebefore the development of a full sum-
mer canopy. It is restricted to sites where the light
intenslty does not fall below 10 per cent of the day-
light between April and mid-June, in which p e r l o d
the flowers are produced. Shoots expand during the
late winter and early spring. The seeds are gradually
shed,mainlyinJulyandAugust.The leaves are
normallydead by July. There is then a period of
aestivation (dormancy during the dry season). This
ends in the autumn when a new set of roots forms.
The plant cannot replace damaged leaves and is very
foliage contains toxlc glycosides and, though sheep
and cattle will eat it, rabblts will not. Its reproduc-
tive strategy is intermediate between a stress-tolerant
ruderal and a competltor-stress-tolerator-ruderal (p.
103). It extends to 340 m around Sheffield, but is
known to grow up to 660 m in the British Isles. It IS
largely absent from skeletal habitats and steep slopes.
The bluebell commonly occurs in woodland. In the
Sheffieldsurvey, I t was recordedmostfrequently
rn broad-leaved plantations. It was also common in
scrub and woodland overlying either acidic or lime-
stone beds, but less frequentinconiferousplanta-
tions. It occurs in upland areas on waste ground and
heaths, and occasionally in unproductive pastures, on
spoil heaps, and on cliffs. In woodland habitats, it
40 LIFE AND THE ENVIRONMENT

70 - Phanerophytes

60 -
1 Chamaephytes
Hernicryptophytes
Cryptophytes
A
Y Therophytes
C
a,
50-
a,
Q
v
v)

E
0
40-
L

-"
-
'i; 30-
5
.-Y0
5 20-
&
10 -

0-
Tundra
Normal spectrum Tropical Temperate Desert

Figure 2.10 The proportion of plant life-forms in various ecozones. The 'normal' spectrum was constructed by selecting
a thousand species at random.
Source: From data in Raunklaer (1934) and Dansereau (1957)

growsmorefrequentlyand is significantlymore constrained by limiting factors In their environment.

abundanton
south-facing
slopes.
However,in Limiting factors Include moisture, heat, and nutrient
unshaded habitats, I t prefers north-facing slopes. It is
levels. Each species hasa characteristic tolerance range
not found in wetlands. It can grow on a wide range and ecological valency. Limiting climatic factors are
of soils, but I t most frequent and more abundant in radiation and light, various measures of temperature
the pH range 3.5-7.5. It is most frequent and abun- (e.g. annual mean, annual range,Occurrence of frost),
dantinhabitatswithmuchtreelitterandlittle various measures of the water balance (e.g. annual
exposed soil, though itis widely distributedacross all precip~tation,effective precipitat~on, droughtp e r l o d ,
bare-soil classes. snow cover), windiness, humidity, and many others.
Of these climatic factors, temperature and water are
master limiting factors. They are summarized in cli-
SUMMARY mate diagrams that characterize bioclimates. Ecozones
are large climatic regions sharing the same kind of
All living things live in partlcular places - habnats. climate. The Mediterranean ecozone is an example.
Habitats rangeIn size from few a cubic centimetres to Ecozones are equivalent to zonobiomes, which in turn
the entire ecosphere. Species differ in thelr habitat are composed of similar biomes. Soil and substrate,
requirements, the span going from habitat generalists, topography, and
disturbancealso
influence life.
who live vmually anywhere, to habitat specialists, Certainsoilsproducedistinctivevegetationtypes
who arevery choosy about their domicile. Speciesare called pedobiomes, such as that associated with soils
 LIFE A N D THE ENVIRONMENT 41

Altitude 30 i Slope Aspect

301 m
C
a,
3 20 n.@
0
0
0
L

10 E 10
Unshaded
z
n
L

2
0 0
0 100 200 300 400 500 0 20 40 60 80 100
Metres Degrees from horlzontal

56.8
P< 0.001

Per cent abundance

Hydrology

0 0 ' e100
0 '.-j!{
9 5 0 Water
depth (mm)
"
1
" 1
1
-cuLou)
I I -

.f
Per cent bare soil
* Ma
gn1to open water
a

Figure 2. I I Autoecological accounts for the bluebell (Hyucinthordes non-xnptu) In the Sheffield reglon, England. For the
subject species, bluebell in the example, the blank bars show the percentage of s~mpleoccurrences over each of the
environmental classes,and the shaded bars show the percentage of samples mwh~ch20 per cent or more ofthe quadrat
subsect~onscontained rooted shoots. The hangular ordination' is explained in Figure 4.8.On the 'aspect' diagram,
n.s. stands for 'not significant'.
Source: After Grime et 4l. (1988)
42 L I F E AND T H E ENVIRONMENT

formedon serpentme. Several topographic features FURTHER READING

Influence life - altitude (which produces orobiomes),
aspect, inclination (which along a hillslope produces
vegetation catenae), and insularity (the relatlve ISO-
latlon of a community or ecosystem). Disturbance by
bioticandabiotic factors is an importantenviron-
mental factor. Examples are disturbance by wmd and
fire. Life has to adapt to environmental conditlons in
the ecosphere. There are several 'ways of living', each
ofwhlch corresponds to an ecological nlche. Ecological
equivalents (or vicars)are species from different
stock, and livlng In different parts of the world, that
have adapted to the same envlronmental constralnts.
Adaptation to environmental conditlons is also seen
In life-forms. The overall habltat preferences of an
individual species requlre detailed and intense study.
They may be summarlzed as autoecologlcal accounts.
Cox, C. B. and Moore, P. D. (1993) Biogeography: A n
Ecological and Evolutlonavy Approach, 5th edn, Oxford:
Blackwell.
Dansereau, P. (1957) Biogeography: A n Ecological
Perspectizle, New York: Ronald Press.
Forman, R . T. T. (1995) Land Mosaza: The Ecology of
Lands~zpes and Regions, Cambridge: Cambrdge
Unlverslty Press.
Huggett,R. J . (1995) Geoerdngy: A n Ez~olrrtronavy
Appmdrh, London: Routledge.
Kormondy, E. J . (1996) Concepts of Erology, 4th edn,
Englewood Cliffs, NJ: Prentice Hall.

ESSAY QUESTIONS Larcher, W . (1995)Edogy: Physrologd

Plant
Ecophysiology and S t w s Pbyslolngy of Frrnctlonal Groups,
1 How useful are bioclimatic edn,
3rd Berlin: Sprmger.
classifications to ecological
biogeographers?
Stoutjesdijk,
and
P.
Barkman, J. J. (1992)
2 How important is disturbance
in Microdimate. Vegetation and Fauna, Uppsala, Sweden:
communities?
shaping Opulus Press.

3 Why are ecological

equivalents so Viles, H . A. (ed.) (1988) Biogeomorphology, Oxford:
common?
Basil Blackwell.

Whelan, R. J. (1995) The Ecology ofFire, Cambridge:

Cambridge University Press.
 3
THE DISTRIBUTION OF O R G A N I S M S
All species a n d other gronps of organism have a particular geographical range or distrihution.
This chapter couers:
kinds of distribution
howorganismsspread
how species distributions are broken apart
human impacts on species distributions
C O S M O P O L I T A N AND PAROCHIAL:
P A T T E R N S OF D I S T R I B U T I O N
All species, genera, families, and so forth have a geo-
graphical range or distribution. Distributions range
in size from a few square metres to almost the entire
terrestrlalglobe.Theirboundarles are determined
by the physical environment, by the living envron-
ment, and by history. They tend to follow a few basic
patterns - large or small, widespread or restricted,
continuous or broken.
large and small, widespread and
restricted
An endemic species lives in onlyone place, no
matter how large or smallthat place should be. A
species can be endemic to Australia or endemic to a
few square metres in a Romanian cave (when it is a
micro-endemic).A pandemic species lives inall
places. The puma or cougar ( F e h concoIor), for exam-
ple, is a pandemic species because it occupies nearly
all the western New World, from Canada to Tierra
del Fuego (Colour plate 2). It I S also an endemic
species of this region because it is foundnowhere
else. Cosmopolitan species inhabit the whole world,
though not necessarily in all places. I t is possible for
a cosmopolitan specles to occur ~n .numerous small
localities in all continents. As arule,pandemic or
cosmopolitan species have widespread distributions,
whereas endemic species have restricted distribu-
tions. The Romanian hamster (hIesocricetus newtmi) is
restricted andendemic(Figure 3.1). The capybara
(HydrotSoerus hydrochaerzs), the largest living rodent,
is endemic to South America. It is also a pandemic,
ranging over half thecontinent.Thedistinction
betweenwidespread and restrictedoftenrestson
the occurrence or non-occurrence of species within
continents or biogeographical regions (Table 3.1).
Micro-endemic species
Some species have anextremelyrestricted (micro-
endemic) distribution, living as a single population
in a small area. The Devil’s Hole pupfish (Cyprznodon
dzabolis) is restrictedtoonethermalspringissuing
from amountainside in south-westNevada (Moyle
andWilliams 1990). The Amargosa vole (hlicrot/ds
cillifornirus srzrperrszs) lives inasingle watershed In
California, Unlted States (Murphy and Freas 1988).
The rare black hairstreak butterfly (Strymonidzd p r ~ n i )
is confined to a few sites in central England and In
central and eastern Europe.
44 THE DISTRIBUTION OF ORGANISMS
Figure 3.1 The Romanlan hamster ( M e s m m z u ~newtoni), a specles wlth an endemlc and restricted distribution. It lives In
a narrow strlp along the Black Sea coast In Bulgaria, Romanla, and adjacent parts of the Ukraine.
Source: After Bjarvall and Ullstrom (1986)
Endemic plant families
Two restricted and endemic flowering-plant families
are the Degenertaceae and the Leltneriaceae (Figure
3.2). The Degenerlaceaeconsists of asingletree
spectes, Degenma uztzenszs, that grows on the Islandof
Fiji. The Leitnerlaceae also consistsof a single species
- the Florldacorkwood (Lertnerzcz flWidZna). This
dectduousshrubisnatlvetoswampy areas in the
south-eastern Untted States where t t is used as floats
for fishing nets.
Cosmopolitan plant families
Twowidespreadflowenng-plantfamiliesarethe
Mesocricetus newtoni
sunflowerfamily(Compositae or Asreraceae) and
the grass family (Graminae) (Figure 3.3). The Com-
positaeisone of thelargestfamilies of flowering
plants. I t c o n t a m around 1,000 genera and 25,000
species. It is found everywhere except the Antarctic
mainland, though it is poorly represented in troplcal
rainforests. The grass family comprises about 650
genera and 9,000 species, including all the world’s
cereal
crops (including rice). Itsdistributtonis
world-wide, ranging from inside the polar cmle to
the equator. It is the chlef component in about one-
fifth of the worlds vegetatlon. Few plant formattons
lack grasses; some (steppe, pratrte, and savannah) are
dominated by them.
 THE DISTRIBUTION OF O R G A N I S M S 45

Table 3. I Speciesclassed according to range size and cosmopolitanism

Range of cosmopolitanism

Endemic (peculiar) Characteristic Semicosmopolitan Cosmopolitan

(shared between (shared between (shared between
two three or four five or more
biogeographical biogeographical biogeographical
regions) regions) regions)

Microscale hairstreak Skua

Black Friesea (Stercorarius Stenotypic
butterfly oligorhopala - a skua) - a coastal ornamental plants
(Strymonidia prund collembolan bird of Antarctica,
Amargosa vole species found in southern
South
(Microtus Tripoli (Libya), America,
Iceland,
californicus Malta, Bahia andFaeroes
the
scirpensis) Blanca
Devil’s Hole (Argentina), and
pupfish Santiago (Chile)
(Cyprinidon
diabolis)

Mesoscale California vole Rose pelican Olivaceus warbler Cormorant

(Microtus (Pelecanus (Hippolais pallida) (Phalacrocorax
californicus) onocrotalus) - a passerine bird carbo) - a bird
Romanianhamster - a bird shared shared b southern from the
(Mesocricetus
newton!)
by central Asia t:
Europe, t e Near Palaearctic,
and southern East, and scattered Oriental,
Africa places in the Ethiopian,
Ethiopian region Australian, and
Nearctic regions

Macro- and Capybara Puma or cougar Black heron Human (Homo

megascale
(Hydrochoerus (Felis concolor) (Nycticorax sapiens) - a
hydrochaeris) - a pandemic of nycticorax) - a pandemic
- a pandemic the New World pandemic bird of cosmopolite
South America,
North America,
Africa, and Eurasia

Source: After Rapoport (1 982)

Zonal climatic distributions shrub specles, I S pantroplcal, though centred In the

Some animal and plant distributlons follow climatic
zones (Figure 3.4). Five relatively common zonal
patterns are pantropical (throughoutthetropics),
amphitropical (eltherslde of the tropics), boreal
(northern), austral (southern), and temperate (mid-
dle
latitude).
The sweetsop and soursop family
(Annonaceae), conslstmg of about 2,000 trees and
Old World tropics. Thesugarbeet,beetroot,and
spinachfamily(Chenopodiaceae)consists of about
1,500 species, largely of perennial herbs, widely dis-
tributed elther side of the tropics in saline habitats.
The arrowgrassfamily (Scheuchzeriaceae) comprises
a singlegenus (Srherrrhzevza) of marsh plantsthat
are restricted to a cold north temperate belt, and are
especially common In cold Sphagnrm bogs. The
 THE DISTRIBUTION OF ORGANISMS 47

Figure 3.4 Zonal climatic distributlons of four plant families: the pantroplcal sweetsop and soursop family (Annonaceae),
theamphitropical sugarbeet, beetroot, andspmachfamily(Chenopodiaceae),the cold temperatearrowgrassfamily
(Scheuchzerlaceae), and the north temperate poppy family (Papaveraceae).
Source: After Heywood (1978)

Jump dispersal disjunctions
Jump dispersal IS the rapldpassage of Individual
organisms acrosslargedistances,oftenacross
hospitableterrain. Thejump takeslesstlmethan
the individuals'life-spans.Plantsandanimals
survive a long-distance jump and founda new colony
groups spread slowly along mountain chains. Later,
membersoccupylngthecentre of the distributlon
became extmct, leaving the surviving memberson
in- elther side of the tropics.
Humans carry specles to all cornersof the globe. In
that doing so, they create disjunct distributions. A prime
example 1s the mammals Introduced to New Zealand.

lead to 'jump' disjunctions. The process is probably There are fifty-four suchIntroducedspecies (C.M.
common, especially in plants. There are about 160 King 1990). Twenty came directly
or indirectly from
temperate or cool temperate plant specles or species Britam and Europe, fourteen from Australia, ten from
groups that have amphltroplcal distributions in the the Americas, SIX from Asla, two from Polynesla, and
Americas(Raven1963).Most of thesearosefrom two from Africa. The package contalned domestic ani-
Jump dispersal. Exceptions are membersof the wood- mals for farming and household pets, and feral animals
landgenera Osmmbna and Sanrcula (Raven1963). for sport or fur production. Farm anlmals Included
Troplcalmontane specles of thesegeneraalmost sheep, cattle, and horses. Domestic animals included
bridge the gap in the dislunctlon. This suggests that cats and dogs. Sporting animals Included pheasant,
thedisjunctions areevolutlonaryinorigln. The deer, wallabies, and rabbits. The Australian possum
48 THE DISTRIBUTION OF ORGANISMS

was introduced to start afur industry. Wild boar and for the disjunct distributlonof the flightless runnmg
goatswereliberatedonNewZealandbyCaptain birds, or ratites (p. 7 1).
JamesCook.Manyother specles wereintroduced
- Europeanblackblrds,thrushes,sparrows, rooks,
yellowhammers, chaffinches, budgerigars, hedge- Climatic disjunctions
hogs, hares, weasels, stoats, ferrets, rats, mlce. Several
Climatic disjunctionsresult from a once widespread
speciesfailed to establishthemselves.Thesefailed
distribution
beingreduced
and
fragmented by
antipodeansettlersmcludebandicoots,kangaroos,
climaticchange.Anexample IS themagnoliaand
racoons, squirrels, bharals, gnus, camels, and zebras.
tulip treefamily(Magnoliaceae)(Figure 3.5). This
family conststs of about twelve genera and about220
Geological disjunctions tree and shrub specles natwe to Asia and America.
The three American genera (Magnolia, Talauma,and
Geological disjunctions are common In the south-
Lznodmdron) alsooccur In Asla.Fossil formsshow
erncontinents,whichformedaslngleland mass
that the family was once much more wldely distrib-
(Gondwana) during the Trlassicperiod but have sub-
utedintheNorthernHemisphere,extendinginto
sequentlyfragmentedanddriftedapart.Ancestral
GreenlandandEurope.Indeed,theMagnoliaceae
populatlons livmg on Gondwana were thus split and
wereformerlypart of anextensiveArcto-Ternary
evolved independently. Thelr Gondwanan orlgln is
declduousforest
that covered much Northern
reflected In present day distributions. Thls IS seen in
Hemlsphere land until the end of the Tertiary period,
many flowering plant families. The protea, banksia,
whenthedeclineintothe Ice Agecreatedcold
and grevillea family (Proteaceae) IS one of the most
climates.
prominentfamilies In theSouthernHemlsphere
(Figure 3.5). It provides numerous examples of past
connectlons between South Amencan, South Afncan, Relict groups
and
Australian floras. Thegenus Gevuina, for
instance, has three species, of whlch one is native to These are produced by envlronmental changes, par-
Chileandtheother two toQueenslandandNew tlcularly climatlc changes, and evolutionaryprocesses
Guinea. The breakup of Pangaea is also responsible that lead to a shrmking distribution.

Figure 3.5 The protea, banksia, and grevillea family (Proteaceae), and the magnolia and tulip tree family (Magnoliaceae).
The Proteaceae or~g~nated on Gondwana and have survivedon the all southern contments. The Magnoliaceae once formed
part of extenswe Northern Hemisphere declduous forests that retreated from hlgh latitudes durlng the Quaternary Ice
age.
Sourre: After Heywood (1978)
 THE DISTRIBUTION OF ORGANISMS 49

Climatic relicts widespread In northern and central Europe. Climatic

warmingduringtheHoloceneepoch hasleft it
Climaticrelicts aresurvivors of organismsthat stranded in Norway, the Ural Mountains, and two
formerly had larger distributions. The alpine marmot isolated sites in Scotland.
(Mamota mamota), a large ground squirrel, lives on
alpine meadows and steep rocky slopes in the Alps
(Figure 3.6). It hasalsobeenintroducedinto the Evolutionary relicts
Pyrenees, the Carpathians,and theBlackForest.
During the IceAge it livedontheplainsarea of Evolutionary relicts aresurvivors of ancient groups
central Europe. With Holocene warmlng, it became of organism. The tuatara (Sphenodon punctatus) is the
restricted to hlgher elevations and Its present distri- only native New Zealand reptile (Plate 3.1). It is the
butlon, which lies between 1,000 m and 2,500 m, is sole survivingmemberthereptilianorderRhyn-
a relict of a once much wider species distribution. cocephalia, and is a 'living fossil'. It lives nowhere
The Norweglan mugwort (Artemisia n m g z c a ) is a else in the world. Why has the tuatara survived on
small alpine plant. During the last Ice age, it was NewZealandbutotherreptiles(andmarsupials

Figure 3.6 The alpme marmot (Murmotu murmotu) - a climatlc relict.

Source: After Bjarvall and Ullstrom (1986)
50 THE DISTRIBUTION OF ORGANISMS

Plate 3.1 Tuatara (Sphenodon punc)phrs)

Photograph by Pat Morris

andmonotremes) have not?NewZealandinthe widely distributed

during
the Mesozoic
era -
Cretaceous period lay at latitude 60" to 70" S. The specimens have been unearthed in Oregon, Green-
climatewould have beencolder,and thewinter land,Siberla,andAustralia.They may have been
nlghts longer. Most reptiles and mammals could not popular items on dinosaurian menus. The reduction
havetoleratedthisclimate.Thetuatara has alow of cycad distribution slnce the Mesozoic may have
metabolic rate, remaining actwe at a temperature of partlyresultedfromcompetitionwithflowering
1l0C, which is too cold to allow activity in any other plants(whichreproducemoreefficientlyandgrow
reptile. It may have survived because it can endure faster). Climatic change may also have played a role.
cold conditions. Its long isolation on an island, free Overthelast 65 million years,tropicalclimates
from competltron with other reptiles and mammals, slowly pulled back to the equatorial regions as the
may also have helped. worldclimatesunderwentacooling. Cycads are
Cycads belongtothefamily Cycadaceae, which therefore In partclimaticandinpartevolutionary
comprises nine genera and about a hundred species, relicts. They commonly maintaln a foothold in iso-
all of whlch are very rare and have highly restricted lated reglons - their seeds survive prolonged imrner-
distributlons confined to the tropics and subtropics sion in sea water and the group has colonized many
(Colour plate 3). Early members of the cycads were Pacific Islands.
 THE DISTRIBUTION OF O R G A N I S M S 51

Range size and shape east-west range dimenslons for North American snake
species(Brown 1995).Inthegraph, ranges equi-
Interesting relationships distant In north-south and east-west directions will
The areas occupied by species vary enormously.In lie on the line of equality thatslopes at 45 degrees up-
Central and North America, the average area occu- wards from the origin (Figure 3.7b). Ranges stretched
out in a north-south direction will lie below the line
pied by bear species (Ursidae) is 11.406 million km';
of equality; ranges that are stretched in an east-west
for cat species (Felidae) it is 5.772 million km'; for
squirrel, chipmunk, marmot, and prairie dog specles direction lie above the line of equality. The pattern
(Sciuridae) I t IS 0.972 million km'; and for pocket for North American snakes is fairly plain (Figure
gopher species (Geomyidae) i t is 0.284 million km' 3.7a). Small ranges lie mainly abovethe line (theseare
(Rapoport1982:7).The rangeoccupied by indi- stretched in a north-south direction); largeranges
vidual species spans100 km' for such rodents as tend to lie below the line (these are stretched in an
thepocket mouse Heteronlys dtsmaresttant~s to 20.59 east-west direction).
million km' for the wolf (Canis Iuprt~). There is a plausiblereason for these patterns, which
Species rangein CentralandNorth America is are also found in lizards, birds, and mammals (Brown
related to feeding habits (Table 3.2).Large carnivores 1995: 110-11). Species with smallgeographical
tend to occupy the largestranges. Large herbivores ranges are limited by local or regional environmental
come next, followed by smaller mammals. The larger conditions. The major mountain ranges, river valleys,
range of carnivores occurs in African species, too. The and coastlines in North America run roughly north to
mean species range for carnivores (Canidae, Felidae, south. The soils, climates, and vegetation assoclated
andHyaenidae) is 8.851million km'; the mean with these north-south physlcal geographical features
specles rangefor herbwores (Bovidae, Equidae, Rhino- may determine the boundariesof small-range species.
cerotldae, and Elephantldae) I S 3.734 million km'. Large-range species are distributed over much of the
continent. Theirranges cannot therefore be influenced
by local and regional environmental factors. Instead,
Geographical regularities in ranges they may be limited by large-scale climatic and vege-
The size and shapeof ranges are related. Figure 3.7ais tational patterns,which display a zonary arrangement,
ascattergraph of thegreatest north-south and the running east-west in wide latitudinal belts.

Table 3.2 Mean geographical ranges of Central and North American mammal species grouped
by order

Order Mean range area

(millions of km2]

Carnivora(carnivores) 6.174
Artiodactyla (even-toed ungulates) 5.072
Lagomorpha (rabbits, hares, and pikas) 1.926
Chiroptera (bats) 1.487
Marsupialia (marsupials) 1.130
lnsectivora(insectivores) 1 . 1 17
Xenarthra or Edentata (anteaters, sloths, and armadillos) 0.889
Rodentia (rodents) 0.764
Primates (primates) 0.249

Source: After Rapoport (1 982)

52 T H E D I S T R I B U T I O N OF ORGANISMS

U T
0
I I
2,000
I I
4,000
East-west distance (krn)
I
6,000 0 2,000
-4,000
East-west distance (km)
7
00

F t p m 3.7 (a) A scattergraph of the greatest north-south versus the greatest east-west dimensmn of North Amerlcan
snake specles ranges. The line represents ranges whose two dimensions are equal. (h) Some example shapes and snes of
ranges. For convenience, the ranges are shown as squares and rectangles, hut they could be clrcles, ellipses, or any other
shape. Thediagonallinerepresenrsrangeswirhtwoequaldimensions.Abovetheline,rangesarestretchedin a
north-south directlon; helow the line, they are stretched in and east-west directlon. Sor/rw: Afrer Brown (1995)

Range size tends to increase with increasing Distribution within a geographical

latitude.Inother words,onaverage, geographical
ranges are smaller In the troplcs than they are near the
poles. Moreover, spec~eswhose ranges are centred at
increaslng hlgher latitudes (nearer the poles) tend to
be distributed over anincreasinglywiderrange of
latitudes. This relationship is called Rapoport’s rule
(Stevens 1989), after its discoverer (see Rapoport
1982).The same pattern holds for altltudinaldis-
tributlons: wlthin the same latitude, the altltudinal
range of a spec~esincreases with the midpolnt eleva-
tlon of the range(Stevens 1992). These b q e o g r a -
phicd patterns are baslc, holding for organlsms from
land mammalsto coniferoustrees. Five hypotheses
may explaln them - continental geometry, congenial
envlronments,constraints
on dispersal, climatlc
change,and
troplcal
competltlon (Brown1905:
112-14) (Box 3.1).
Rapoport’srule does notapply everywhere. The
range size of Australian mammals does not increase
from the troplcs towards the poles (F. D. M. Smith
et a/. 1994). Latitudinal and longltudinal variatlons
in range size correlate wlth continental wldth. More-
over, the arid centre of Australia contains the fewest
mammal species withthe largestranges, whilethe
molst andmountainous east coast, andthemon-
soonal north, contain large numbers of specles with
small ranges.
range
Home range
Indivlduals of a specles, especially vertebrate species,
may have a home range. A home range is the area
traversed by an individual (or by a pair, or by a family
group, or by a social group) in its normal activities of
gathering food, mating, and caring for young (Burt
1943). Home ranges may have irregular shapes and
may partly overlap, although individuals of the same
speclesoften occupy separate home ranges. Land
Iguanas (ConolophnJ pallid~~s) on the island of Sante
Fe, In the Gakpagos Islands, have partly overlapping
home ranges near cliffs andalongplateauxedges
(Figure 3 . 8 4 (Christlan et al. 1983). Red foxes (Vzdpes
zmlpes) in theUnlversity of Wisconsmarboretum
have overlapping ranges,too (Figure 3.8b) (Ables
1969).
The sue of home ranges varies enormously (see
Vaughan 1978: 306). In mammals,smallhome
rangesareheld by the female prairie vole (Microtus
ochrogaster), mountain beaver (Aplodontia rufi), and
the common shrew (Sorex armem), at 0.014 ha, 0.75
ha, and 1.74 ha, respectlvely. The American badger
(Tuxideu taxzrs) has a medium-size home range of 8.5
km’. A female grizzly bear (Urws avstos) with three
 THE DISTRIBUTION OF ORGANISMS 53

Box 3.1
ACCOUNTING FOR REGULARITIES and persistence of narrowly endemic species. The
IN R A N G E S I Z E logic of this ideais that a species living at high
latitudescould pass over mountainsinsummel
Continental geometry withoutexperiencingharsherconditionsthan it
experiencesnormally.In low latitudes, however
Rapoport’s rule and the tendency of small ranges to speciesused toliving intropicallowlandsanc
occur at low latitudes follow from the geometry of trying to cross a mountain pass atanequivalenl
NorthAmerica.Thecontinenttapersfromnorth elevation would experience conditionsnever experi.
tosouth so speciesbecomemoretightlypacked enced before, no matter what the season. The samc
in lower latitudes. Against this idea, species ranges would be true of a tropical montane species trying
become
smaller
before the
tapering becomes to cross tropical lowlands in the winter.
marked. Moreover, it doesnotaccount for the
elevational version of Rapoport’s rule.
Climatic change
Congenial environments Large-range species have adapted to the Pleistocenc
climatic swings at high latitudes.
The abiotic environments in low latitudes are more
favourableand lessvariable than thoseinhigh
latitudes.Small-rangespeciesliving in low lati- Tropical competition
tudes canavoidextinction by survivingin‘sink’ Range size increases and species diversity decrease
habitats, and recolonize favourable ‘source’ habitats inmovingfromtheequatortowardsthe poles
and re-establish populations after local extinctions. Interactionbetweenspecies may be astronge
limiting factor in lower latitudes.
Constraints on dispersal
Barriers to dispersal
are
more
severe
in
low
latitudes,which may contribute to the evolution

yearlings has a large home range of 203 km’, and a (Vestzarzu coccmea), a Hawaiian honeycreeper, exhibits
pack of elght tlmber wolves (Canis lupus) has a very terrltorlal behaviour when food is scarce.
large home range of 1,400 km2.
Habitat selection
Territory
As the conditlonsnear the margms of ecological toler-
Some anlmals actlvely defend a part of thelr home ancecreatestress, I t followsthatthegeographlcal
range against membersof the same species. This core range of a specles IS strongly influenced by Its eco-
area, whlch does not normally lnclude the perlpheral logical tolerances. It IS generally true that specles with
parts of thehomerange, is called theterritory. wide ecologlcal tolerances are the most wldely dis-
Species thatdividegeographicalspaceinthis way tributed. A specles will occupy a habltat that meets its
areterrltorlalspeaes.Terrltorlesmay be occupied tolerance requlrements,for I t slmply could not survwe
permanently or temporarily.Breeding p a m of the elsewhere. Nonetheless, even where a population is
tawny owl(Strzx aluco) stay within their own territory large and healthy, not all favourable habltat Inside Its
during adulthood. Great tlts (Parus m a j o r ) establish geographical range will necessarily be occupled, and
terrltorles only during the breeding season. The ‘i‘iwi theremay beareas outsideItsgeographicalrange
54 THE DISTRIBUTION OF ORGANISMS

Cliff

7-
100 m

(b) -"
.""""" -.
%
A .-.-..
Female 1
Female 2
8
I
I
*
I

"""".
I
I

-
I
Female 3 I
I
II
I

**
I

1 km Male 1 I
I
8 m
t I Male 2 8 m
I

Figure 3.8 Home ranges.(a) Land iguanas .(ConoIophuc pulliduc) on the island of SanteFe, in the Gal6pagosIslands.
(b) Red fox (Vd@ wul@) home ranges in the University of Wisconsin arboretum.
Sources: (a) After Christian et ul. 1983; (b) After Ables (1969)
 THE DISTRIBUTION OF ORGANISMS 55

where I t could live. In many cases, indivlduals 'choose'confined to areas where the mean annual temperature
notto live In particularhabitats, a process called of the coldest month exceeds -O.S"C, and,like
habitat selection. Not a great deal is known about madder, seems unable to withstand low temperatures
habttat selection, but it doesoccur andlimitsthe (Iversen 1944). Several frost-sensitlve plant species,
distribution of some species. including the Irish heath (Eriru erigena), St Dabeoc's
Habitat selection is prevalentin birds.An early heath (Daboecra cantabrim), large-flowered butterwort
study was carrled out in the Breckland of East (Prngmxla grandifla), and sharp rush (J/INLXS acutw),
Anglia, England (Lack 1933). Thewheatear (Oenanthe occuronlyin the extreme west of the British Isles
ornunthe) lives on open heathland in Britain. It nests where winter temperatures are highest. Otherspecles,
in old rabbitburrows. I t does not occurin newly such as the twinflower (Lintmu boredis) and chick-
forested heathland lacking
rabbits.
Nestmg-slte weed-wintergreen (Trientalis rtrropaea),have a northern
selection thus excludes it from otherwlsesuitable or north-eastern distribution, possibly because they
habitat.Thetreeplplt (Anthm tvrvra1i.c) andthe have a winter chilling requirement for germmation
meadow plpit (Anthus pratetuis) are bothground that southerly latitudes cannot provlde (Perring and
nesters and feed on the same variety of organisms, Walters1962). Low summertemperatures seem to
but the tree pipit breedsonly In areas with one or restrict the distribution of such species as the stemless
more tall trees. In consequence, in many treeless areas thistle (Cirsmrn ac-a/&). Neartoitsnorthernlimit,
in Brltain, the tree pipit I S not found alongside the thls plant I S found malnly on south-fmng slopes, for
meadow piplt. David Lack found some tree pipits on north-faclng slopes it fails to set seed (Pigott 1974).
breeding in one treeless area close to a telegraph pole. Thedistribution of grey halr-grass (Corynepphorm
The pipits used the pole merely as a perch on whlch czznemns) I S limited by the 15°C mean isotherm for
to land at the end of their aerial song. Meadow pipits July. Thls may be because itsshort life-span (2to
singaslmilarsongbut landon theground.This 6 years) means that, to maintain a population, seed
finding suggests that the tree piplt does not colonize production and
germinationmustcontinueun-
heathland slmply because I t likes a perch from which hampered (J. K. Marshall 1978). At the northern limit
toslng.The concluslon of the Breckland study of grey halr-grass, summertemperatures arelow,
was that the heathland and pme planation birds had whichdelaysflowering, and, by the time seeds are
a smaller distributionthan they otherwisemight produced,shadetemperatures are low enoughto
because they selected habitat to live in. In short, they retard germinatlon.
were choosy about where they lived. The small-leaved lime tree (Tilia cordatu) ranges
across much of Europe. Its northern limit in England
and Scandinavla IS marked by the mean July 19°C
Distributional limits
isotherm(Pigott1981;PigottandHuntley1981).
Thetree requires 2,000growing day-degrees to
Plants
produce seeds by sexual reproduction.But for the
Many distributional boundaries of plant species seem lime tree to reproduce, the flowers must develop and
to result from extremeclimaticeventscausingthe then the pollen must germmate and be transferred
failure of one stage of the life cycle (Grace 1987). The through a pollen tube to the ovary for fertilization.
climatic events in question may occur rarely, say once Thepollen fails togerminateattemperaturesat
or twice a century, so the chances of observing a fail- or below 15"C,germinates best intherange17°C
ure are slim. Nonetheless, edgesof plant distributions to 22OC, and germinates, but less successfully up to
often coincide with isolines of climatic variables. The about 35°C. A complicating factor is that the growth
northern limit of madder (Rubia pmgrina) in north- of thepollentubedependsontemperature.The
ern Europe sits on the 40°F (4.4"C) mean January growth rate is maximal around 20-25OC, diminish-
isotherm (Salisbury 1926). Holly (Ilex aquifolium) is ing fast athigherand lower temperatures.Indeed,
56 THE DISTRIBUTION OF ORGANISMS

the extenslon of the pollen tube becomes rapid above ature; mean length of the frost-free permd; potentla1
19"C,which suggest why thenorthernlimit vegetation; mean annual precipitatlon; average gen-
is
marked by the 19°C mean July isotherm. eral humidity; and elevatlon.Isolines for average
Several models use known climatlc constramts on mlnimum January temperature, mean length of the
plant physiology to predict plantspecies distributlon.frost-free perlod, and potential vegetation correlated
One studyInvestigated the climatlcresponse of boreal with the northern range limits of about 60 per cent,
tree species In North America (Lenihan 1993).Several 50 per cent,and 64 per cent, respectively, of the
climatic predictor variables were used In a regressionwlntering bird spec~es. Figure3.1 1 shows the wlnter
model.The variables were annual snowfall, day- distributionandabundance of the easternphoebe
degrees, absolute minlmum temperature, annual (Suyornisphoebe). The northernboundary is constrained
soil-moisture deficlt, andactualevapotranspiration by the -4'C isotherm of January minimum temper-
summed over summermonths.Predictedpatterns ature.Justtwoenvlronrnental factors - potential
of species' dominanceprobability closely matched vegeration and mean annual precipitation - coinclded
observed patterns (Figure 3.9). The results suggested with eastern range boundaries, for about 63 per cent
thatthe boreal tree species respond individually and 40 per cent of the species, respectively. On the
to different comblnations of climaticconstraints. western front, mean annual precipitatlon distributlon
Another study used a climatlc model to predict the colncided for 34 per cent of the species, potential
distribution of woody plant specles in Florida, Unitedvegetatlon 46 per cent, and elevation 40 per cent.
States (Box et ul. 1993). The State of Florida is small Why shouldthenorthernboundary of so many
wlnterlng bird specles coinclde wlththe
enough for variatlons in substrate to play a malor role average
in determining what grows where. Nonetheless, the minlmum January temperature? The answer to this
model predictedthatclimatic poser appears to lie in metabolic rates (T. L. Root
factors, particularly
winter temperatures, exert a powerful Influence, and 198th). At their northern boundary, the calculated
in some cases adirectcontrol,on mean metabolic rate in a sample of fourteen out of
specles' distri-
butions. Predicted distributions andobserved distrib- fifty-one passerine (song birds and thelr allies) specles
ution of the longleaf pine (PZNWpulustris) and the was 2.49 times greater than the basal metabolic rate
(which would occur in the thermal neutral zone, see
Florida polson tree ( M e t o p z m toxzjierm) are shown in
Figure 3.10. Thepredictions for the longleaf pine p. 20). This figure implies that the w m e r ranges of
are very good, except for a narrow strip near the cen-thesefourteen bird species are restricted to areas
tralAtlantlc coast. Thematch between predicted where the energy needed to compensate for a colder
and observed distributions is not s o good for the environment I S not greater than around 2.5 times the
Florlda polson tree. The poison tree is a subtropical basal metabolic rate. The estlmated mean metabolic
species and the model was less good at predicting the rate for thirty-six of theremamingthlrty-seven
distribution of subtropical plants. passerine species averaged about 2.5. Thls '2.5 rule'
applies to birds whose body weight ranges from 5 g
in wrens to 448 g in crows, whose diets range from
Animals
seeds to insects, and whose northernlimlts range
Many bird species distributions are constrained from Florida to Canada - a remarkable finding.
by such environmental factors as food abundance,
climate,habitat,andcompetltlon.Distributions of
148 species of North American land blrds wintering SPREADING: DISPERSAL AND
In the conterminous Unites States and Canada, when RANGECHANGE
compared with environmentalfactors, reveal a consis-
tent pattern (T. L. Root 1988a, b). Six envlronmental Some organisms - information is too scanty to say
factors were used: average minimum January temper- how many - have an actual geographical range that
 THE DISTRIBUTION OF ORGANISMS 57
Predicted boreal torest

\$$
Picea-Pinus
Picea-Popu/us-Pinus
..........
. .. . . . . .. : .. I.
..............
.............. Picea-Abies
f-l

Picea-Pinus-Acer-Abies

Observed boreal forest

Figure 3.9 Boreal forest types In Canada. (a) Predicted forest types using a regression model. (b) Observed forest types.
Sortm: After Lenihan (1993)
58 THE DISTRIBUTION OF ORGANISMS

1
Lu

.I.
LI

Predicted**coo'll' .'

8.""- - - -" - ", -..,

Florida poison tree

Metopium toxiferum

Figure 3.10 Predictedandobservedlongleaf pine (Pinus palustrn) and the Florida polson tree (Metoprum t o x f m m )
distributlons in Florida, United Stares.
Source: After Box et al. (1993)

is smaller than t h e n potentla1 geographical range. In Jump dispersal IS the rapid transit of individual
other words, some species do not occupy all places organisms across large
distances,
often
across
that their ecological tolerances would allow. Often,a inhospitableterrain.Thejumptakes less time
species has simply failed to reach the 'missmg bits'. thanthelife-span of theindividualinvolved.
Dispersal IS the process whereby organisms colonlze Insects carrted over sea by the wtnd IS an example
new areas. It is a vast subject that has long occupled (see p. 60).
the minds of ecologists and biogeographers. Diffusion is the relatively gradualspread of
populations acrosshospitableterram. It takes
place over many generatlons. Specles that expand
Life on the move then ranges little by little are said tobe diffusing.
Organismsdisperse.They do so In at leastthree Examples Include the Amerlcan muskrat(Ondatra
different ways (Pielou 1979: 243): zi6ethzrm), spreading in central Europe after five
 THE DISTRIBUTION OF ORGANISMS 59

-.-.I
.-.-.__
,

Figure 3.1 I Winter distribution andabundance of the eastern phoebe (Suywnu phoebe). The northern boundary 1s
constrained by the 4°C average minimum January Isotherm.
Soum: Map after T. L. Root (1988a, b); picture from Stokes and Stokes (1996)

individuals were introduced by Bohemlan

a Agents of dispersal
landowner in 1905 (Elton 1958). It now inhabits
Europe In many millions. Species may disperse by active movement (digging,
3Secularmigration is the spread or shift of a flying, walking,or swimming), or by passive carriage.
species that takes place very slowly, so slowly that Passive dispersal IS achieved with the aid of physical
the species undergoes evolutlonary change whileit agencies (wind, water) or biological agencies (other
is takingplace. By the time population arrives in a organisms, Including humans). These various modes
new region I t will differ from the ancestral popula-of transport are given technical names- anemochore
tlon in the source area. South American members for wind dispersal, thalassochore for sea dispersal,
of the familyCamelidae(thecamelfamily) - hydrochore for water dispersal, anemohydrochore
the llama (Lama glama), vicufia (Lama uzcugna), for a mixture of wind and water dispersal, and bio-
guanaco (Lama guanrroe), and alpaca (Lama pacos) chore for hitching a ride on other organisms. Figure
- are examples. They are all descended from now 3.12 shows the means by which colonists were carried
extinct North American ancestors that underwent to Rakata, which lies in the Krakatau Island group,
a secular migration during Pliocene times over the after the volcanic explosion of 1883. Notice thatsea-
then newly created Isthmusof Panama. dispersed (thalassochore) species, most of which live
60 T H E D I S T R I B U T I O N OF ORGANISMS

125 Phase I Phase

fleece (it was difficult to search all parts of the anlmal),
1 ; 2 produced 8,5 11 seeds from 85 specles. The seeds were
a mlxture of hooked, bristled and smooth forms. They
included sweet vernal-grass (Anthoxumthrrtn odorutmz),
100
large thyme (Thynus pdegioides), common rock-rose
lady's bedstraw (Gulirlrn
(Heliunthen11mn11711t)11~lar/ri?~l),
vernm), and salad burnet (Summorbu nunor).
.I 75
01
P
c
0 Good and bad dispersers
8
.n
5
z 50
Dispersal abilities vary enormously. Thls is evldent
in records of thewidest ocean gap known to have
been crossed by varlousland animals, either by fly-
ing,swimming, or on rafts of soil andvegetation
25
(Figure 3.13). The 'premier league' of transoceanrc
dispersers I S occupled by bats and land blrds, insects
and splders, andland molluscs. Lizards, tortoises, and
0 rodents come next, followed by small carnlvores. The
poorest dispersers are large mammals and freshwater
fish.
Figure -3. I ? The means by which the spermatophyte flora Not alllarge mammals are necessarily ineptat
reached Rakata, In the Krakarau Island group, from 1x83 crossing water. It pays to check their swlmming pro-
to 1989. The collarlon periods are Indicated o n the
ficiency before drawingtoo manybiogeographlcal
horlzontal axis. Human introduced species are excluded.
S w m r : After Bush and Whitcaker (1991)
conclusions from their distributions. Fossil elephants,
mostlypygmy forms, are found on manyislands:
along strand-lines, are rapid colonizers. The anemo- San Miguel, Santa Rosa, and Santa Cruz, all off the
chores comprisethree ecological groups.The very Californiancoast;Miyako andOkinawa,both off
early colonists are mostly ferns, grasses, and compos- China; Sardinia, Sicily, Malta, Delos, Naxos, Serifos,
ltes (membersof the Compositae),whlch are common Tilos, Rhodes,Crete,
and
Cyprus, all in the
in early pioneerhabitats. Asecond group is domlnated Mediterranean Sea; and Wrangel Island, off Siberia.
by forest ferns, orchids,and Asclepiadaceae (milk- Before reportsonthe proficiency of elephants as
weed, butterfly flower, and wax plant family). These swlmmers (D. L. Johnson 1980), it was widely
second-phase colonists requlre more humidcon- assumed thatelephantsmust have walked to these
ditions. Numerically, mostof them areepiphytes. The islandsfrom mainland areas, taklngadvantage of
thlrd groupconsists ofseven prlmarily wlnd-dispersed former land brldges (though vicarianceeventsare
trees. Anlmal-dispersed (zoochore) organisms are the also a possibility). Now it is known that elephants
slowest to colonlze. They are malnly carried by birds could, apparently, have swum tothe islands, new
and bats. explanations for the colonization of the islandsare
The efficacy of organlsms as agents of dispersal I S required.
surprising. A study carrled out in the Schwabische Supertramps are ace dispersers. They move with
Alb, south-west Germany, showsjust how effective ease across ocean water and reproduce very rapidly.
sheep are at spreading populations of wild plants by They were first recognized on the island of Long, off
dispersing their seeds (Fischer et al. 1996). A sheep New Guinea (Diamond 1974). Long was devegetated
was specially tamed to stand still while I t was groomed and defaunated about two centuries agoby a volcanlc
for seeds. Sixteen searches, each covering half of the explosion. The diversity of bird species is now far
 THE DISTRIBUTION OF ORGANISMS 61

‘large‘ mammals

A Freshwater
fish
r I 1 I I I I 1
0 500 1,000 1,500 2,000 2,500 3,000 3,500
Distance (km)
pigum 3.13 The widest mean gaps crossed by terrestrial anmals. The distances are extremes and probably not cYPlcal of
the groups.
Sourte: After Gorrnan (1979)

higher than would be expected (Figure 3.14). Of the Dispersal routes

forty-three species present on the island, nine were
responsible for thehighdensity.Thesewerethe The ease andrate at whichorganlsmsdisperse
supertramps. They specialize in occupymg islands too depend on two things: the topography and climate
smalltomaintainstable,long-lastingpopulations, of the terrain over which they are moving and the
or islands devastated by catastrophic disturbance - wanderlust of a particular species. Topography and
volcanic eruptions, tsunamis, or hurricanes. They are climate mayimpose constraintsupon dispersmg
eventually ousted from these islands by competitors organlsms. Obviously, dispersal will be more easily
that can exploit resources more efficiently and that accomplished overhospitable terrainthanoverin-
can survwe at lowerresource abundances. hospitable terrain. Obstacles or barriers to dispersal
may be classified according to the ‘level of difficulty’
in crossing them. George Gaylord Simpson (1940)
suggested three types:
62 T H E D I S T R I B U T I O N OF O R G A N I S M S

1 Stablelandbridges.These are‘filterroutes’in
Long Island Simpson’s sense. Faunas are free to move in both
directions.
2 Periodicallyinterruptedlandbridges.These
are akin to stable land bridges, but there are two
differences. First, access in both directions is pen-
odically interrupted by water gaps. Second, faunas
on one or both sldes may suffer extinction, owing
to the loss of area during tlmes of separation.
3 ‘Noah’s arks’. These are fragments of lithospheric
plates carryingentire faunas withthem from
one source area to another (see p. 76). Ordinarily,
transport on Noah’s arks I S one-way.
0 40 80 120
Number of species present 4 Stepping stone islands. These are a falrly
permanent or temporary serles of islands separated
F&re 3.14 Birdabundanceonvariouslslandsbetween by moderate to small water gaps. Traffic could be
New Gulnra and New Brltain. Abundance was measured
by thedailycapturerate In nets. Nectlng ylelds,which
two-way, but oftengoesfromislands of greater
reflectcoral populariondensltles, increase with rhelocal diversity to Islands of lesser diversity.
number ofspecles.LongIsland is exceptlonal because 5 Oceanicislands.These are situated a long way
populatlon denslty I S abnormally high for an island of I ~ S from the mainland and they receive ‘waifs’. They
size. This hlgh density IS due largely to the presence of
are similartosteppingstone islands, butthere
superrramp species.
S o r m ~ e :Afrer Diamond ( 1 974)
will be fewer arrivals that arrive at much longer
time intervals. They are nottrue sweepstakes
routes because the chances of arriving depends on
1 ‘Level 1 ’ barriersare corridors - routes through
species characteristics - some species have a much
hospitableterrain
that allow theunhindered
better chance of arrwing than others.
passage of animals or plants in both directions.
‘Level 2’ barriers are filter routes. An example is
Of course, for terrestrial animals, crossing land is not
a land bridgecombinedwith a climatlc barrier
so difficult as crossing water, and many large mam-
that bars the passage of some migrants. ThePana-
mals have dispersed between biogeographical regions.
manian Isthmus is such a route since it filters out
There I S probably no barrler that cannot be crossed
species that cannot tolerate tropical conditions.
given enough time:
‘Level 3’ barriersare sweepstakes routes. They
reflect the fact that, as in gambling,there are Onemorning[inGlacierPark],darkstreaks were
always a small number of wlnners compared wlth observed extendingdownward at variousanglesfrom
saddles or gaps in the mountains to the east of us. Later
losers. In biology, the winners are those few lucky
In theday,thesestreaksappeared to be muchlonger
individuals that manage to survive a chance jour- and at the lower end of each there could be discerned
ney by water or by alr and succeed in colonizing adarkspeck. Through binocularsthese spots were
places far from their homeland. seen to be animals floundering downward in the deep,
softsnow. As theyreachedlowerlevelsnor so far
Simpson’s terms apply primarily to
connections distant,theyproved to be porcupines.From every
between continents. A morerecent scheme, though little gap there poured forth a dozen or twenty, or in
not dissimilar to Simpson’s, is more applicableto one case actually fifty five, of these animals, wallowing
down to thetimberlineonthe west slde.Hundreds
connections between Islands andcontinentsand
of porcuplneswerecrossingthemainrange of the
island with other islands (E. E. Williams 1989). It Rockies.
recognlzes five types of connections: ( W . T. Cox 1936: 219)

Dispersal today
Dlspersal undoubtedly occurs at present but I t I S nor-
mallydifficult to observe. The chiefproblem In
detecting dispersal in actlon is that detailed species
distributions are scarce. Most instances of organisms
movlng to new areas probably pass unnoticed - is a
new sightlng an Individual that has movedinfrom
elsewhere, o r is I t anIndividualthat was born In
the area but has not been seen before! In addition, it
is sometimes difficult to account for a range change.
Species ranges alter
through dispersal
and local
extinctlons. Actlng In tandem, dispersal and extinc-
tlon may It.ad t o range expansion (through all or
any of the dispersal processes), to range contraction
(from local extlnction), o r to range 'creep' (through
a mlxtureofspreadand local extlnction).It is bar
from easy t o establish the processes Involved In actual
cases.
Despite the problems of studylng range changes,
there are several amazlng cases of present-day disper-
sal resulting from human Introductions. Introduced
species are taken to new areas, accidentally or p u r -
posely, by people. Not allintroductlonssurvlve;
somegaln a foothold but progresslittlefilrther;
others go rampant and swiftly colonize large tractsof
what is to them uncharted territory. Just four species
of amphibians and reptiles live In Ireland, compared
with twelve on the British mainland. The species are
the natterjack toad (B//fildutuitu), the common newt
(Trit//ru.r ~ w / g u r i ~the
) , common or viviparous lizard
(Lucwtu z'rzllpm-u), and the common frog ( R u m tempo-
vuriu). The common frog was introduced into ditches
In University Park, Trinity College Dublin, in 1696,
Itstillflourshestheretoday,and has spread to
the rest of Ireland. So, why have only three species
of amphibiansandreptilescolonized Ireland! One
explanation is that other newts and toads did estab-
lish bridgeheads, but they died outbecause they were
unable to sustain large enough colonles for successful
Invasion. Thls view is lentsomesupport by the
present distributlon of the natterjack toad In Ireland
- I t I S restricted to a small part of Kerry and shows
no slgns of sprading.
Afine example of a rampant disperser I S the
THE DISTRIBUTION OF ORGANISMS 63
Europeanstarling (Strrvmrs d g a r i s ) . Thisbird has
successfully colonized North America, South Africa,
Australia,and NewZealand.ItsspreadinNorth
Amerlca was an indubltable ecologicalexplosion -
within sixty years it had colonized the entire United
States and much of Canada. There were several 'false
starts' or failed introductions during the nineteenth
century when attempts to introduce the bird in the
UnitedStatesfailed.Forexample, In 1899, twenty
pam of starlings were released in Portland, Oregon,
but they vanished. Then in April 1890, eighty birds
were released in CentralPark,NewYork,andin
March the following year a further eighty birds were
released. Within ten years the European starling was
firmly established in the New York City area. From
that staglng post, I t expanded its range very rapidly,
colonlzlng some
7,000,000km' in fifty years
(Figure 3.15). The speed of dispersal was due to the
Irregular migrations and wanderingsof non-breeding
1 - and 2-year-old starlings. Adult birds normally use
thesamebreedingground year after year anddo
notcolonizenewareas. The roamlngyoungbirds
frequented faraway places. Only after five to twenty
years of migration between the established breeding
groundsandthe newsites, didthebirdstakeup
permanent residence and set u p new breeding
colonles. For example, the European starling was first
reported in California In 1942. It first nested there in
1949. Large-scale nestingdidnotoccuruntilafter
1958.
Dispersal in the past
Centres-of-origin and dispersal
Historical biogeography began as a scholarly debate
about the restocking of the Earth from Mount Ararat
after Noah's Flood. The debate evolved into the clas-
sical theory of centres-of-originanddispersal first
proposed by Charles Robert
Darwinand
Alfred
Russel Wallace. Darwln argued that all species have
a centre of orlgln from which they disperse; and that,
because barriersimpedetheirmovement, specles
spread slowly enough for naturalselection to cause
them to change while dispersing.H e saw dispersal as
64 THE DISTRIBUTION OF O R G A N I S M S

t -O0\ x I

Figure 3. I5 The westward spread of the European starling (Sturnus vulgtzrrs) In North Amenca.
Source: Map after Kessel (1953) and Perrins (1990); plcture from Saunders (1889)

a phenomenon of overarchmgimportance,andhis last 65 million or so. O n two occasions during

centre-of-origin-disprsal model became the ruling
theory In historical biogeography. It was developed
persuasively in thetwentieth century by Ernst Mayr,
George Gaylord Simpson, and Philip J. Darlington.
South American invasions
A classic example of a dispersal model is the mam-
malian invasions of South America (Simpson 1980).
Although South Amerlca IS now connected to North
America, I t was an island-contment for most of the
that time, a land connection with North Amenca,
probably through a chain of islands, developed for a
few million years and was then lost. Durmg these
times, and in recent times, mammals Invaded South
America from the north. Four phases of invasion are
recognized (L. G. Marshall 1981a) (Figure 3.16):
1 Phase I, whichoccurred In the LateCretaceous
per& andearliestPalaeoceneepoch, was an
Invasion by ‘old timers’ or ‘ancient immigrants’.
Three orders of mammal invaded - only two of

. .
60 50 40 30
Millions of years ago
Figure 3.16 T h e history of South American mammals.
L. G. Marshall (1981a)
Sourre: After
which are represented in South Amerlca today
the marsupials, the xenarthrans(edentatesin
some classlficatlons), andthe condylarthrans. Once
isolated
again,
these
ancestral
stocks
evolved
independently of mammals elsewhere and some
unique forms arose. The first marsupial invaders
were didelphoids and included ancestors of the
modernopossums (Box3.2). The borhyaenlds
weredog-likemarsupialcarnivores.Theybore
a strong resemblance to the Australian thylacine
or Tasmanian wolf, the likeness being the resultof
convergentevolution. Thyfacosrnifus, amarsupial
equwalent of the sabre-toothed tigers, was also a
splendid exampleof convergent evolution (Figure
3.18). The xenarthrans(‘strange-jointedmam-
mals’) were ancestors of living armadillos, sloths, 3 Phase I11 began in the Late Miocene epoch, some
andanteaters,andextinctglyptodonts(Figure
3.19). The condylarthrans produced a spectacular
array of endemic orders of hoofed mammals, all
of whicharenowextinct - the condylarthra,
litopterns,noroungulates,astrapotheres, tngon-
stylopoidea, pyrotheres, and xenungulates (Figure
3.20).
THE DISTRIBUTION OF ORGANISMS 65
Phase IIIb
Camelidae, Canidae, Cervidae,
Equldae, Felidae, Leporidae,
Gomphotheriidae, Mustelidae,
Sciuridae, Soricidae, Tapiridae,
Tayassuidae, Ursidae
Phase IIIa
Crlcetidae, Procyonldae
Phase I1
Prlmates, Cavlomorph rodents
..
Phase I
..... Astrapotheria, Condylarthra,
.. Litopterna, Manuplalia,
. ... Notoungulata, Xenungulata,
Pyrotherla, Tngonostylopoldea,
Xenarthra (Edentata)
20 10 0
- 2 Phase I1 Involved abrief‘window of dispersal
opportunity’ arising in the Late Eocene and Early
Oligocene epochs, from about 40 million to 36
million years ago. Duringthistrrne,aseriesof
islands linked the two American continents. Two
groups of mammalinvadedSouthAmerica -
primates and ancestorsof the caviomorph rodents.
Thesewerethe‘ancientislandhoppers’.After
having a r m e d In SouthAmerica,bothgroups
underwentanImpressiveadaptiveradiationto
producethegreatvarlety of rodentsfoundin
South America today (aswell as some interesting
extinctforms - the LatePleistocene Teficwnys
gigantissirnus was nearly as large as a rhinoceros)
and the New World monkeys.
6 million yearsago. Then,theBolivarTrough
connectedtheCaribbean Sea withthe Pacific
Ocean and deterred thepassage of animals (Figure
3.21).PhaseIIIa saw the first mammals rafting
across the seaway on clumps of soil and vegeta-
tion.These‘ancientmarlners’weremembers of
two families - the ‘field mouse’ family (Crlcetidae)
66 THE D I S T R I B U T I O N OF ORGANISMS

V " Y

fish, and frogs (Figure 3.17). It is the only aquatic marsupial in the world. The female's pouch is closed
by a sphincter muscle when diving to prevent her babies from drowning.

and the racoon and its allies family (Procyonldae).
By 3 million years ago,there was acomplete
land connection - the Panamanian land bridge -
that furnished a gateway for faunal interchange
betweenNorthandSouthAmerica.PhaseIIIb
began as a flood of mammals simply walked into
South America. Members of many families were
involved - camels(Camelidae),dogs(Canidae),
deer (Cervidae),horses (Equidae), cats (Felidae),
rabbits (Leporidae), mastodons (Gomphotheri-
idae), weasels (Mustelidae), squirrels (Sciuridae),
shrews (Soricidae), mice (Muridae), tapirs (Tapir-
~dae),peccaries (Tayassuidae), and bears (Ursidae).
The traffic was two-way - many South American
species travellednorthwardsandenteredNorth
America.
4 Phase IV startedabout 20,000 years ago(the
exactdate is debatable) as humansspreadinto
South America.
The outcome of these waves of invasion is distinct
faunal strata inthepresentmammals of South
America. Anteaters, sloths, armadillos, and opossums
are survivors of the first invasion. The caviomorph
 THE DISTRIBUTION OF ORGANISMS 67

Smilodon Thylacosmilus
Placental sabre-tooth Marsupial sabre-tooth

Figtm 3.18 Convergent evolutlon of the rnanuplal sabre-tooth Tby/acormifusand the placental sabre-tooth Smilodon.
Source: After L. G. Marshall (1981a)

rodents and New World monkeys are survivors of the tral taxa had been fragmented by tectonlc, sea-level,
second Invasion. All other South Amerlcan mammals and climatlc changes. He termed the fragmentation
(saverecentintroductions)aresurvivorstheGreat process‘vlcariism’ or ‘vlcariant
form-maklng In
Amerlcan Interchange. immobilism’. Fortunately, it has becomeknown as
vicariance. Of course, to become wldespread in the
first place,aspecies must disperse.Butvicariance
SPLITTING: VICARIANCE AND
biogeographersclaimthatancestraltaxa achreve
RANGECHANGE
widespread distributlon through a mobile phase zn
the ubsenre of buwzers. They allow that some dispersal
What is vicariance?
acrossbarriersdoesoccur, but feel that I t is a
Vicariance blogeography arose as an antldote to the relatively insignificant blogeographical process.
hegemony ofdispersal biogeography. Antl-dispersalist
grumblings were heardearly In the twentieth century.
Continental fission
The first major crltlcal onslaught was directed by U o n
Croizat, a Franco-Italian scholar (Croizat 1958,1964). Continents break up, drift, and collide. In doing so,
Croizattested theDarwiniancentre-of-origin- theymakeandbreakdispersalroutesandalter
dispersalmodel by mappingthedistributlons of species distributrons. Overall, continental drift, and
hundreds of plant and anlmal species. H e found that processes associated with it (such as mountain build-
species wlthqulte differentdispersalpropensities ing), help to explain several features in animal and
andcolonizingabilitieshadthesamepattern of plant distributions for both living and fossil forms.
geographicaldistribution.Hetermedtheseshared The tearing asunder of previously adjoining land
geographlcaldistributionsgeneralized or standard masses causes the separation of ancestral populatlons
tracks. Crolzat argued that standard tracks do not of animals and plants. Once parted, the populations
represent lines of migration. Rather, they are the pre- evolve independently and diverge. Eventually, they
sent distributions of a set of ancestral distributlons, may become quite distinct from the ancestral popu-
or a biota of which Individual components are relict lation that existed beforethe land masses broke apart.
fragments. His reasoning was that widespread ances- There are several good examples of this process.
68 THE DISTRIBUTION OF O R G A N I S M S

Figure 3.19 A glyptodont. Thls tanklike Pletstocene herbwore was descended from old xenarthran Invaders.
Sourre: After L. G. Marshall (1981a)

i Litopternsh
”

Condylarths

Pyrotheres

Astrapotheres Notoungulates

Figure 3.20 Unique South Amerlcan mammals that evolved from Late Cretaceous condylarthran Invaders.
Source: After Simpson (1980)
 THE DISTRIBUTION OF ORGANISMS 69

Before 3 million years ago After 3 million years ago

\

Figure 3.21 The Bolivar Trough and Panamanian Isthmus. Before 3 million years ago, a marine barrier separated Central
America and South Amenca. When the barrierdried up, the PanamanIan land brldge was formed. Animals ventured
northwards and southwards In the Great Amerlcan Interchange.
Sourre: After L. G. Marshall (1981b)

Reptiles and mammal-like reptiles on freshwaters, it is difficult to imagine it breasting

Pangaea the South Atlantlcwaves for 3,000 miles’.
Anotherdisjunctdistribution is displayed by
Thebreakup of Pangaea,andinparticularthe Lystrosaurus, a squat, powerful, mammal-like reptile.
southern partof it knownas Gondwana, split several Lystrosaurus lived in the Early Trlass~cperlod, about
animal and plant populations into separate groups. In245 million yearsago. It was about a metre long,
consequence, members of these ancient populations, with a palr of downwards-pointing tusks. The posi-
andtheirlivingdescendants, have dis~unctdistri- tion of its eyes, high on Its head, suggest that 1t spent

butions. Oneof the first piecesof fossil evrdence used much tlme submerged with all but the top of its
1n support of the idea that Africa and South Amerlca head below water. Specimens of Lystrosaurus have
were formerly ~ornedwas the distributlon of a small been found at localities 1n India, Antarctica, South
reptile called Mesosaurus that lived about 270 million Africa (all formerly part of Gondwana), and China
yearsago,In thePermianperiod(Figure 3.22). (Figure 3.23). Thedistribution of Lystrosaurus on
Mesosaurus was about a metre long, slimly built with Gondwana accords with modern ranges of terrestrial
slender limbs and paddle-like feet. At the front end reptiles,such as thesnappingturtle, Chelydra ser-
it had extended, slender laws carrymg very long and pentrna (Colbert 1971). Its presenceinChina,ina
sharpteeth,probablyusedtocatch fish or small place that IS normally assumed to have been part of

crustaceans. The rear end was a long and deep tail, eastern Laurasia in Triassic times, presents a problem.
admlrably suited to swrmmrng. It surely spent much One explanation, thoughanunlikelyone,isthat
time in thewater. The sedimentsIn which it has been some individuals migratedfromGondwana, all
found suggest that it inhabited freshwater lakes and the way round the head of the Tethys embayment,
ponds. Its remalns have been found only in southern toeastern Laurasia. Anotherexplanationisthat
Brazil and southern Afnca. If, as the evldence sug- Gondwana was biggerthan is normallysupposed,
gests, it were a good swimmer, then it would have and extended beyond the northern edgeof the Indian
had a widerrangethanitapparently did have. plate to include a large segment of what is now China
Themostparsimoniousexplanation 1s that Brazil (Crawford 1974). If thiswerethe case, allthe
andAfrlcaabuttedintheLate Palaeozoicera. As Lystrosaurus faunas would have lived on Gondwana,
Alfred Sherwood Romer (1966: 117) said, ‘although and the Chinese members of the population would
Mesosaurus was obviously a competent swlmmer In not be anomalous. Another possibility, suggested by
70 THE D I S T R I B U T I O N OF O R G A N I S M S
T South
Americ
jd
Figure 3.22 The distribution of Merosaurus in the Permlan period
workonplatetectonm In theSouth-eastAsian
region, is that In Late Permian times, or even before,
large fragments of Australia (terranes) appear tohave
broken off anddriftednorthwards,collidingwith
Asia. These terranes could have carried, in the man-
ner of Noah's arks, Ly~trosaunrswith them, or could
have acted as a serles of steppingstonesmaking
dispersal possible (C. B. Cox 1990: 125).
Cenozoic land mammals
During Cenozoictimes,land-animalevolutlon was
greatly affected by contmental drift (Kurt& 1969).
Fossil Cenozoic faunas, and particularly the mammal
faunas, are fairly differentoneachcontinent.The
most distinctive faunas are those of South America,
Africa, andAustralia - thesoutherncontinents.
There are some thirty orders of mammal, almost two-
thirds of which are alive today. All the mammalian
orders probably had a common orlgln - an ancestor
that lived in the Mesozoic era. The Cenozoic diver-
gence of mammals may be due to the Late Mesozoic
fragmentation of Pangaea.Interestingly,theland-
dwelling animals (mostly reptiles) that lived in the
Cenozoic era displayed far less divergence than the
k
Africa
mammals. By the end of the Cretaceous p e n o d , after
about 75 million years of evolution, some seven to
thirteenorders of reptileshadappeared, far fewer
than the rhirty or so produced by mammalian evo-
lution over 65 million years. A possible explanatlon
for thlsdifferenceliesinthepalaeogeography of
the Mesozoic contlnents. For much of the Age of
Reptiles, the continents were not greatly fragmented.
There were two supercontinents: Laurasia lay to the
northandGondwanatothesouth.Riftsbetween
the continents existed as early as the Triassic period,
but they were not large enough to act as barriers to
dispersaluntilwellintotheCretaceousperiod.In
Late.Cretaceous and Early Tertiary times, when the
mammals began to diversify, the rapid breakup of the
formerPangaea,coupledwithhighsea-levels,led
to the separation of several land masses and genetic
isolation of animal populations. The result was diver-
gence of early mammalian populations.
Pangaea and plant distributions
The breakup of Pangaea accounts for many disjunct
plant distributlons. Some plants have seeds unsuit-
ableforjumpdispersal.Anexampleisthegenus
 71

t
Figwe 3.23 The distribution of Ly~tm~uww In the Early Triassic period.
Soume: Base map after A. G. Smith et al. (1994)

Nothofugw (thesouthernbeeches)thatconslsts of Big, flightless birds

about sixty species of evergreen and deciduous trees
and shrubs. Its present distributionis disjunct, being Continental breakup helps to explain the enigma of
found on remnants of Gondwana - South America, the large flightless blrds. This group forms an avian
New Zealand, Australia, New Caledonia, and New superorder - the ratites. There are five families of
Guinea, but not Africa.Fossil Nothofugxr pollen of living ratites and two extmct ones, all of which are
OligoceneagehasbeenfoundonAntarctica.The thought to have arisen early in bird evolution from
conclusion normally drawn is that the modern dis- acommonancestorandthusareamonophyletic
junctrangeofthegenushasresultedfromthe group.Themoas(Dinornithidae)livedonNew
breakup of Gondwana. The same explanation applies Zealanduntila few hundred yearsago; the kiwis
to plants of the protea, banksia, and grevillea family (Apterigidae) still live on New Zealand (Plate 3.2).
- the Proteaceae (p. 48). Emus(Dromaiidae)andcassowaries(Casuariidae)
72 THE DISTRIBUTION OF ORGANISMS
Plate 3.2 Brown or spotted kiwi ( A p b x australis)
Photograph by Pat Morris
both live in Australia, and the cassowariesarealso
found on New Guinea (Colour plate 4). The extinct
dromorthinlds are also Australian ratltes. Ostriches
(Struthlonidae) live in Africa and Europe. The ele-
phantblrds(Aepyornlthidae)livedon Madagascar
and possibly Afrlca (e.g. Senutet al. 1995), but went
extinct around the mid-seventeenth century. Rheas
(Rheidae) live in South Amenca.
With the exception of the chlcken-sizedkiwis,
most of the ratites are well-built cursorial blrds wlth
huge hind limbs for fast and sustamed running. The
ostrich IS the world’slargestlivingbird,standing
about 2.5 m high and weighing In at about 140 kg.
The largest of the moas, Dzornrs maxmus, stood about
3.5 mhighandweighedaround 240 kg; i t was
the tallestbirdknowntohavelived. The largest
elephant-bird, Aepyornis maximus, was a ponderous
glant, wlth elephantlne-style legs; I t stood 3 m tall
and weighed about 450 kg. How could such large
blrds have spread through the southern contments if
they could not fly, or even if they could? A plausible
hypothesls involves the early evolution of flightless-
ness and contmental drift (Cracraft 1973, 1974). A
flyingancestor of all theratitesprobablylivedin
west Gondwana, or what is now South Amerlca. The
flightlesstinamous,partridge-likebirdsthat have
thelr own superorder (the Tinamae), maybe descen-
dants of this extinct bird that stayed at home in South
America. Flightlessness may have evolved during the
Cretaceous period, before continental separation was
faradvanced. The flightlessbirdscouldthen have
dispersed through the southern continents by walk-
ing to other partsof Gondwana. The ancestorsof the
NewZealand moas andkiwismust have walked
 THE DISTRIBUTION OF ORGANISMS 73

through west Antarctlca before New Zealand drifted NorthandSouth Amerlca wasfilledby the lower
away from the m a n land mass. Interestingly, a fossil central Amerlcan land mass. However, the malnland
ratite has recently been discovered in the Palaeogene relatives
of Nesophonte.r and Solenodon on North
La MesetaFormation,SeymourIsland,Antarctlca Amerlca were by now extlnct, leaving thelr Antillean
(Tambussi et ui. 1994). Therecently extinct elephant relatlves as relicts of a once wldespread distributlon.
birds of Madagascarcould have walkeddirectly The history of the Greater Antillean insectivores
from South America. The emus and cassowarles may may beinterpreted in other ways. Thetraditional
have walked through east Antarctica. The ostrlches view is that they colonlzed the islands by over-water
of Africa andEurope may have walkeddirectly dispersal from the Americas - a sweepstakes route.
from South America, and the South American rheas Support for this interpretation came from a study of
presumably evolved from less adventurousSouth Caribbeantectonics, thecomposition of thefauna,
American flightless relatives. and the fossil record in the Americas (Pregill 1981).
The Gondwanan origin of the ratites is challenged The evidence suggested that the Antilles started life
by morerecentwork(e.g. Houde 1988). Forms a volcanic archipelago in the Late Cretaceous epoch.
ancestral to modern ratites appear to have evolved in Modern terrestrlal Vertebrates probably started arrlv-
North America and Europe from the Late Palaeocene ing by over-waterdispersal duringtheOligocene
to the Middle Eocene epochs. This would mean that epoch, when most of the livlng genera In the West
livlngratltesareSouthernHemisphererelictsofa Indies first appeared, and continued to do s o through
wldespread groupthatprobablyoriginatedinthe the Quaternary. The varlous living genera and spec~es
Early Tertiary period. arenot evenly distributedthroughouttheislands
andthevertebratesarerepresented by remarkably
few orders,families,andgenera. Thlscomposition
Greater Antillean insectivores
is consistentwithanislandblotabuilt up though
Vicarianceevents In the geologicalhistory of the dispersal, rather than vicariance.
Caribbeanregion mayexplain the clistributlon of A
reconstructionof
geolog~cal history in the
relict insectivores In the Greater Antilles. There are Caribbeanreglon may offer acompromlsebetween
twosuchinsectivores - Ne.rophontes and Solmodon. the dispersaland vicarlance cxplanatlonsof the
Nesophonter lived on Cuba, Hispaniola, Puerto Rlco, Antillean insecttvores and other anlmals and plants
the Cayman Islands, and smaller surrounding Islands. (Perfit and Williams 1989). About 130 million years
Soienodon still lives onCubaandHispaniola(Plate ago,theproto-Antilles were a chainof volcanic
3.3). They were, and inSolenodon's case are, shrew-like Islands lylng along a subductlon zone at the Paclfic
anlmals but larger than a normal shrew. SalenoJon I S Ocean rim (Figure 3.24). They stretched 2,000 km
about 15-16 cm and welghs 4 0 4 6 g. It I S nocturnal betweenthe west c o a t of Mexico andthe coast
and lives In caves, burrows, and rotten trees. of Ecuador. Some 100 million years ago, the North
One explanation of the relict insectlvore distribu- Amerlcan and South American plates started tomove
tlon runs as follows (MacFadden 1980). In the Late westwards over the proto-Caribbean sea floor, and the
Cretaceous, NorthAmerlcanandSouthAmerican lslandsdrifted relativelyeastwards, at theleading
land masses were pined by a land mass that was to edge of theCaribbeanplate. By 76 million years
become the Antilles. At thls t m e , ancestral insectl- ago, Cuba struck the Yucatin region of Mexlco. The
vores lived on North
Americaandthe
proto- remaining islands suffered uplift and deformatlon as
Antillean block. Early In theCenozolcera,the they squeezed through the gap between the Yucatbn
proto-Antilles movedeastwards,relatlve to the rest and Colombia. A land brldgc between the Amerlcas
of the Americas, carrylng a cargo of insectlvores wlth (the proto-Costa Rica-Panama land brldgc) had
I t . Later In the Cenozoic era, the proto-Antilles had begun to grow along a new subduction line formed
reached then present position, and the gap between wherethe Pacific Ocean dived
underneath
the
74 THE DISTRIBUTION OF ORGANISMS

Plate 3.3 A relid Antillean insectivore (So/enodon)

Photograph by Pat Morris

Caribbean plate. On occasions, the proto-Antillean which are all small (one from eastern Cuba is at 0.9
islands may have formed a complete dry land con- cmthe smallestfour-legged beast inthe world),
nection between the Americas around this time. The originally came from South America (Hedges 1989).
proto-Antillesstayedclose to the North American Theyappearto have movedalong the
proto-
continent for the next 20 million years, and indeed Antilleanislandchamaround 70 millionto 80
were often connected to it. About 55 million years million years ago. They disembarked when the island
ago, Cuba hit the Bahamas bank, a large limestone reached the Yucatdn, and established a foothold on
platform, and became wedged there. The remaining the North American continent. Today, there are 68
Islands kept movmg eastwards, causing considerable species in Mexlco. The frogs that chose not to dis-
shearagainst the beached Cuba.Theshearstress embark have produced the 139 species found on the
caused the islands to break up and adopt their mod- GreaterAntilleanislandsatthepresenttime.The
ernconfiguration.PuertoRicobroke away from traffic was two-way. While the Eleutberoductylus frogs
Hispaniolaabout35million yearsago;Hispaniola wentashoreontoNorthAmerica, several North
separated from Cuba about 23 million years ago. American species - including pines, butterflies, and
These tectonic changes affected the biogeography todies (a kind of bird) - boarded the islands while
of the area. Frogs of thegenus Eieutberoahtylus, theyweredocked. The presenceofpineson the
 THE DISTRIBUTION OF ORGANISMS 75

76 million years ago I

/
? Cuba strikes
j Bahamas Bank

Start a
. . . . . . Lessen
...... .

3UTH
4ERICP

.... Shallow
sea Arc magnetism
e Island movement
+ Sea floor subduction
- Fault line
+ti+Collision line

Figure 3.24 The geological hlstory of the Caribbean region over the last 100 million yean.
Source: After Perfit and Williams (1989) and Reddish (1996)
76 T H E DISTRIBUTION OF ORGANISMS
Caribbean islands I S difficult to explam by dispersal
because their heavy seeds are not designed to cross
hundreds of kilometres of salt water. Similarly, many
of the butterfliesfoundon theGreaterAntilles,
including the fragile glasswings (family Ithomiidae)
are not good dispersers. Todies are a family of birds
endemic to the Caribbean. They are distantly related
to the kingfishers. They are not powerful fliers. A 30-
million-year-old tody from Wyommg at least allows
thepossibilitythat ancestors of the present birds
could have boarded the islands early in their eastward
travels, perhaps flying theshortdistance across to
Cuba. At same time,ancestors of the primitive msec-
tivores could have crossed over, either by walking or
swimmingtheshortdistance. Most mammals had
not evolved when the Islands were docked against
North America, but ancestors of theseinsectivores
are known from fossils in North America to be at
least 5 5 million to 60 million years old.
Continental fusion
Drifting
continentseventuallycollide
and fuse.
When they doso, they unite toform a new single land
mass. Uplift or a fall of sea-level may also forge a land
bridge between two formerly separated land masses.
Amovingplate may carryspecies (indeed,entire
faunas and floras) over vast distances,actinglikea
glgantic raft or Noah’s ark; i t may also carry a cargo
of fossil forms,and IS like a Viking funeral ship
(McKenna 1973). Thebiotaon these more-or-less
Isolated land masses should attain a steady state In
whlch origination and extmction (turnover) will be
roughly in balance. These biota should be saturated
- all available niches should be filled.
Faunal mixing
Whentwocontinentscollide, or a land brldge is
formed between them, the fauna and flora of the two
continents are free to mix. The mixing process has
four possible outcomes (L. G. Marshall 1981a):
1 Activecompetition occursbetweenspecles or
genera that occupy the same or very similar niches
on the differentland masses. If the native form
should be more efficient at exploitlng resources,
the Invader will be expelled or kept out. Ofcourse,
such unsuccessful invasion attempts are seldom, if
ever, recorded In the fossil record. The alternative
outcome is that an invader is successful and ousts
the native vlcar.
Passivereplacement may occur when, by
chance, a native species sharing a niche with an
invading species happens to go extinct. There is
no competitlon involved - thesurvlvlng species
was merely lucky to be In the rlght place at the
right time.
Intruders may come across a blota with unexplolted
ecologlcal niches and are able to Insinuate them-
selves intothepre-existingcommunitywithout
having an overt affect on it.Theinsinuators
areecologically uniqueand have no discernible
counterparts In the native biota. Insinuating
species will boost the biotic diverslty.
An Invader may have a similar niche to a native
specles and character displacement (p. 118) takes
place so that both species can live together. These
Invaders are competitors-cum-insinuators.
As the faunas and floras mix, so they come toresemble
one another, though they may still retain distinctive
features. The Great American Interchange is perhaps
the mostspectacular and best documented faunal
mixlng event. Itresulted from the fuslon of the North
andSouth American continentsanddemonstrates
many of the possible outcomes of faunal mixing.
The Great American Interchange
The faunas of North and SouthAmerlca began to mlx
at the close of the Miocene epoch, about 6 million
years ago (p. 65). The Great American Interchange
began as a trlckle. It became a flood around 3 million
years ago,when thePanamanianIsthmusformed,
peaked around 2.5 million years ago,and is still
going on at thepresent time.
It was oncewidelybelieved thatthe invasion
of South America by North Amencan specles caused
theextlnction of manynativetaxa. The placental

carnivoresappearedand
outcompeted marsupial
carnlvores,whichbecameextinct. Likewise, South
Amerlcan‘ungulates’ suffered heavylosses incom-
petltlon wlth the northern Invaders. Recent interpre-
tationsoftheevldencepalnt a morecomplicated
picture ofevents. Two cases illustrate thls pomt(L .G.
Marshall 1981a).
Before the Invasions, South American ‘ungulates’
included litopterns (prototheres and macrauchenids)
and notoungulates (toxodons, mesotheres, and hege-
totheres). To these should be added five families of
xenarthrans andeven two of rodents that occupled the
large herbivore niche. Once the Bolivar Trough was
closed, many families of northern ungulates travelled
south(mastodons, horses,tapirs,peccaries,camels,
and deer). There is some evldence that the disappear-
anceofnative‘ungulates’, especially thenotoung-
ulates, began before the appearance of northernrlvals.
The factors causlng this decline were thus not related
to the Interchange. This conclusion is supported by
the fact that no invading northern species were vicars
(ecological equivalents)ofsouthern specles - they
occupiedslightlydifferentniches.Themostlikely
vlcarswere invading ‘camels’ andthecamel-like
macrauchenids. However, these two groups coexlsted
for 2.5 million years.
It was onginally claimed that the South Amerlcan
dog-likeborhyaenlds(marsupials)declined to the
pomt of extinction when In competition with invad-
ing placental carnlvores from the north. However, I t
nowseemsclearthattheborhyaenidswereextlnct
before dogs, cats, and mustelids moved south. Some
large omnivorous borhyaenids declined
and fell
when the waif members of the Procyonidae arrlvedIn
PhaseIIIa(p.65). For instance, Stylocynm, a large,
omnivorous,bear-likeborhyaenid, had a vicar in
Chapa/v/u/anza, a large, bear-like procyonid. In turn,
Chapulrnalaniu vantshed when members of the bear
family (Ursdae) arrived. It I S also possible that the
placentalsabre-tooths(felids) replaced the natlve
marsupial sabre-tooths (thylacosmilids).
WhenthePanamanIanIsthmustriggeredthe
GreatAmericanInterchange, a large ma~orltyof
land-mammalfamilies crossed reciprocallybetween
North and South America around 2.5 million years
THE DISTRIBUTION OF ORGANISMS 77
ago,in Late Pliocenetimes.Initially,there was an
approxlmatebalancebetweennorthward traffic and
southward traffic. During the Quaternary, the inter-
change became decidedlyunbalanced (S. D. Webb
1991). Groups of North American origin continued
to diversify at exponential rates. In North America,
extinctionsmore severelyaffected SouthAmerican
immigrants - six SouthArnerlcanfamilieswere
lostinNorthAmerica,whiletwoNorth American
families were lost In South Amerlca.
R A N G EC H A N G E :H U M A N SA N D
DISPERSAL
To anextent,theactualrange of a specles is a
dynamic, statlstical phenomenon that is constralned
by the environment: in an unchanging habitat, the
geographical range ofa specles can shift owing to the
changing balance between local extinction and local
invasion. And, I t may also enlarge or contract owing
to hlstorical factors, asso plainly shown by the spread
of many introduced species and chance colonizers
In new, but envlronmentally
friendly,
regions.
Humans haveadvertentlyandinadvertentlyaided
and abetted the spread of many species. An example
of a deliberate lntroductlon is the coypu (Myocastov
c o y p ~ ~ swhlch,
), In the 193Os, was brought from South
Amerlca toBritain forIts fur(nutria).Numerous
escapes occurred and it established Itself in two areas:
at a sewage farm near Slough, where a colony lived
from 1940 to 1954, and in East Anglia, with a centre
intheNorfolkBroads (Lever 1979). A concerted
trappingprogramme is believed to have eradicated
thecoypufromBritain(GoslingandBaker1989).
Anaccidentalintroduction was theestablishment
of the ladybird (Chilocoyus nzgvltus) In several Pacific
islands,north-eastBrazil, West Afrlca, andOman
after shipment from other areas (Samways 1989).
The success of many introductions is beyond
question. Indeed, it is ironical that humans are bril-
liantly successful in the unintentional exterminatlon
ofsome specles,mainly throughhabitatalteration
and fragmentation, but hopelessly unsuccessful in the
purposefuleradicationintroducedspeciesthat have
78 T H E D I S T R I B U T I O N OF ORGANISMS
become pests. Invasion success depends on the inter-
action between the invader and the community i t I S
invading. Predicting the fate and impact of a specific
introduced species is very difficult (Lodge 1993). For
instance, domesticandwild-type European rabbits
have been liberated on Islands all over the world
(Flux and Fullagar 1992).Theoutcome of these
lntroductlons ranges from utter failure to rabbit den-
slties so hlgh that the Island is stripped of vegetatlon
and soil.Some rabbltpopulations have survlved
remarkablyadverse conditions for up to a hundred
years and then become extinct.
There are places wherealien anlmalsandplants
are of malor economic and conservation slgnlticance.
Aprimeexample I S New Zealand (Atkinsonand
Cameron1993).Plantintroductlons have averaged
eleven species per year since European settlement in
1840,anddistinctive landscapesare being increas-
ingly altered by weeds. Many Introduced animals act
as disease vectors or threaten native blota.Recent
studies of introduced wasps show adverse effects on
honey-eating and insectivorous birds. Introduced
possums prey on eggs and nestlings of native birds,
damage native forests, andtransmit bovine tuber-
culosis.
A few examples drawn from the animal and fungal
kingdoms will serve to illustrate the rates of spread
andtheimpactthat invaders may have on native
communities.
Animal introductions
American mink and muntjac deer in Britain
The American mink (Mtrstela vison) is amedium-
sized mustelid carnivore withalongbody.It was
introduced to British fur farms from North America
in the1930s. Some individuals escaped and soon
became established in the wild. Mink is now found
inmany parts of theBritainand will continue to
spread (Figure3.25a). As a carnivore, it has had a
rather different impact on n a m e wildlife than other
introduced mammals. A crucial question is whether
themink occupies a previously vacant nichewith
an antlcipatedmild overall ecological Impact, or
whether I t is a species that is endangeringcom-
petltors such as theotter (LutraIutra) and prey
species (including fish stocks).A surveyhelped to
resolve thls issue, anddrew five conclusions (Blrks
1990).First,themink is littlethreattotheotter,
although i t exacerbates otter decline in areas where
theotter is already endangered. Second, themink
probably has, locally, aided the decline of water vole
(Arvicola trrrejtrir). Thlrd, if theminkshould have
hadany effects on waterfowl, then these have not
been translated into widespread population declines.
Fourth, there are grave potential risks of introducmg
mlnkto offshore islands. Fifth,themlnk is not
having a serlous overall Impact on fish stocks, at least
in England and Wales.
Themuntjac deer (M/mtiar/rs reezwi) is small,
standing about 50 cm at the shoulder (Plate3.4). Its
small slze allows I t to live in copses, thickets,
neglected gardens, and even hedgerows (N. Chapman
et a/. 1994). Following the first releases from
Woburn, Bedfordshire, In 1901, the numbersof free-
livlng Reeves' muntjac in Britain remained low until
the 1920s, when populatlons were largely confined to
the woods around Woburn, and possibly also around
Tring in Hertfordshlre. However, in the 1930s and
1940sthere were furtherdeliberateintroductions
inselected areas some distance from Woburn. As a
consequence, the subsequent spread of Reeves' munt-
jac was from several foci (Figure 3.2513). The spread
in the second half of this century has been aided by
furtherdeliberateandaccidental releases, and by
these means new populations continue to established
themselves outside the mainrange. Thus, the natural
spread has been much less impressive than previously
assumed; even in areas with established populations
it takes a long time for muntjac deer to colonize all
the available habitat. The natural rate of spread is
probably about 1 km a year, comparable to that of
other deer species In Brltaln. Arable land classes are
predominantly selected for, andmarginalupland
land classes tend to be avoided. However, long-estab-
lished populations in areas such as Betws-y-Coed in
Wales show thatmuntjac deermaypersist in low
numbers in atypical habltats.

Fi8ut.e 3.25 The distributlon of the Amerlcan mink (Musteh vzson) and muntjac deer (Muntiucur reewsi) in Britain.
Source: AfterH. R. Arnold (1993)
Introduced predators on islands
The Indian mongoose (Hetpestes auropunctatus) is one
of the most potent predators on diurnal ground-for-
aglnglizards.It has beenIntroducedtovarious
islandsworld-wlde In the hope of controllingrats
and other vertebrate pests. Its success in doing so has
been mixed. Its success in reducing and causing the
extrnctlon of native bird and reptile populations is
spectacular (Case and Bolger 1991). Thls IS demon-
strated by a diurnal lizard census on Pacific islands
with and without mongooses (Figure 3.26). Islands
withoutmongooses have nearly a hundredtimes
greaterdiurnallizardabundancethanIslandswlth
mongooses.
THE DISTRIBUTION OF O R G A N I S M S 79
Muntjac deer
Muntiacus reeves1
rn Up to 1959
1960 onwards
Other potent introduced island predators are rats
anddomesticcatsanddogs.Catsandthetreerat
(Rattus rattus) are capable of climbing trees andaffect
specles that the mongoose is less likely to capture.
New Zealand lizards became far less cbmmon after
the mid-nineteenth century, owlng to-cover reduc-
tion through forest clearance and predatlon by cats.
Present-dayislandswithoutmammalianpredators
aroundNewZealandhouseextraordinarilyhlgh
numbers of lizards, wlth one lizard per 3 mr belng
recorded. The same pattern is found elsewhere. High
numbers of diurnallizardsarefoundonrat-free
CousmIsland In the Seychelles andon San Pedro
Martir in the Sea of Cortez, Mexlco. Some lntroduced
reptiles have also caused extmctlons. The Introduced
80 THE DISTRIBUTION OF ORGANISMS

Plate 3.4 Muntiac deer ( M u n f k u s reeves;)

Photograph by Pat Morris

browntreesnake (Bozgatwegularzs) has elimmated ground-foraging emos sktnks(Emoza nigra and Emma
ten blrd specles and severely affected the lizard pop- trossula) are locally extinct, and the two largest skmks
dation on Guam, thelargest of the Mariana Islands. in Fiji have not been seen on these Islands for over a
Introduced predators produce a distinctlve biogeo- century, although they do survive on mongoose-free
graphical pattern, namely a reciprocal c m c u r r e n c e Islands.
pattern. This means that many natlveamphibians
and reptiles occur on Islands wlthout predators, but
A fungal introduction
are absent from Islands wlth predators. An example
IS the tuatara (Sphenodonpunctatus) and Its predator, Around 1900, the Amerlcanchestnut (Castanea
the Polynesian rat (Rattus exrrlans), in New Zealand. dentata) comprised 25 pet cent of the native eastern
On islandswhere the rat is present, the tuatara is hardwoodforest In theUnltedStates.It was a
either absent or not breeding. The largest survivlng valuable forest specles - its hardwood was used for
frog, the Hamilton frog (Lzopelrna hamiltoni), occurs furniture, Its tall stralght timbersfor telegraph poles,
onlyonrat-freeIslands. On Vitl Levu andVanua Its tannm for leather tannmg, and Its nuts for food.
Levu, the two largest Islandsof Fiji, the combmation By 1950, most trees were dead or dymg as a result
of cats and mongooses has proved devastatmg. Two of chestnut blight, a parasitic disease caused by a
 T H E DISTRIBUTION O F O R G A N I S M S 81

500 4 10 Mongooseislands
0 Mongoose-freeislands

a 100

501 0

0
:
0
0

g
cl
0
0
0 0

Native Secondary Clearings, Sugar Suburban

forest forest gardens cane hotels, parks
Highly types Habitat
Undisturbed 4 disturbed

fungal parasite - the sac fungus, Cryphot/ec/r/'l
(Endorhm) purll.r/tmt. The fungus enters the host tree
through a bark wound madeby woodpeckers o r bark-
boringmsects. It growsIntothebarkandouter
sapwood formmg a spreading, oozing sore that cven-
tually encircles o r 'girdles' the truck o r branch. Once
glrdled, the treedies because nutrient supplies to and
fromtheroots are stopped.Thrtlme fromlnitlal
attack to dcath IS two to ten years.
The sac fungus I S native to Asla. In 1913, I t was
found o n Its natural host in Asla - the Chlnese chest-
nut (CmtLtned n / o / / i . ~ r 1 ~ /-~ 7t o) whlch I t does no serlous
harm. It was Introduced Into New York City by accl-
dent, probably In 190.4. on imported Aslan nursery
plants.The first mfected trees were found r n the
Bronx Zoo. The Amencan chestnut I S so susceptible
to CrJphouectriLr that i t wasremovecl from almost Its
entlrcrangewithin fortyyears.All that remalned
were a few Individuals lucky enough t o have escaped
the fungal spores, and adventitlous shoots spouting
from survlvlng root systems.
In the wake of thechestnutblight, several oak
species, beech, hickories, and red maple have become
co-dominants:the oak-chestnut forest has turned
into oak o r oak-hlckory forests. Thls I S evldent in
Watershed /r 1, western North Carolin;l (Figure 3.27).
Here, the disease first attacked In the latc 1920s. The
survey carried o u r In 19.54 shows the orlg1naI forest
composltlon, as the disease hac1 not killed many trees
by that t m e . T h e Amerlcan chestnut IS plainly the
domlnant specles. occupytng over twlce the basal area
of various hlckory species. By 1054, after the forest
hac1 bornethebrunt of thefungalattack,thetree
specles composltIon had altered d r a m a t d l y . T h e
chestnut had all butdisappeared, and hickorws
( C c t r ~ rspp.),
~ thechestnut oak (Qurru/.t pr/t/u.i), the
black oak (Q//ers./o w////im),and the yellow poplar
( L m o ~ / m / r ot/dip+u)
u had become co-hnlnants.
82 THE DISTRIBUTION OF ORGANISMS
Watershed 41, western North Carolina
Despitethe ravages of the chestnutblight,
replantingwith back-crossedtrees
derived from
blight resistant hybrld chestnuts has led to hopes for
a successful replanting programme. Mesic sites (not
too dry, not too wet), especlally shallow coves, with
survlvlng chestnuts may have the greatest potential
for chestnutgrowthandbiocontrol of blight, if
effective hypovirulence, blight resistance, and forest
management measuresaredeveloped. Inaddition,
survlvlng root systems, which occur throughout the
chestnut's natural range, glve hope that a genetically
engineered, less virulent strain of sac fungus might
be released andhelptorestoreavaluableforest
species (Choi and Nuss 1992).
SUMMARY
Species,genera,andfamiliesdisplaythree basic
distributlonal patterns- large or small, widespread
or
1934 1953
restricted, and continuous or broken. Relict groups
areremnants of erstwhilewidespreadgroupsthat
have suffered extinction over much of their former
range,owingtoclimatic or evolutionarychanges.
Rangesizeandshapedisplayrelativelyconsistent
relationships wlth latltude and altmde. Individuals
have home ranges and, sometimes, territories; many
select thehabitatthattheylivein,choosingnot
to occupy all suitable habitat. Distributional limits
to species are determined partlyby ecological factors,
partly by history, and partlyby behaviour. Organisms
disperse. They may do so actively (by walklng, swim-
mlng, flying, or whatever) and passively (carried by
wind,water,orotherorganisms).Dispersalability
varies enormously throughout the living world. Ease
of passage along dispersal routes varies from almost
effortless to nlgh on impossible. Dispersal occurs at
present, much of I t resulting from human introduc-
tions. Dlspersal in the past forms the subject matter
of traditional historical biogeography, where centres-

of-origin and dispersal tootherparts of theglobe
are the prime concern.Vicariance is a rival idea. I t
emphasizes thesplitting of ancestral and widespread
species distributions by geological or ecological
factors. Agoodexample is thebreakup of Pangaea,
which appears to have dictated the current distribu-
tions of several animal and plant groups. The joining
of two or more continents leads to faunal mixing.
A classic example is the Great American Interchange.
E SQSUAEYS T I O N S
1 Why has the centre-of-origin and
provedpopular in
explaining the distribution of
organisms?
2 To what extentpresent
is the
distribution of animal and plant groups
shaped by the breakup Of Pangaea?
3 To what extent have humans
'homogenized' the world biota?
T H E D I S T R I B U T I O N O F ORGANISMS 83
FURTHER READING
Brlggs, J. C. (1995) Globak Btogeography
(Developments in
Palaeontology and Stratigraphy,
14),Amsterdam: Elsevier.
Cox, C. B. and Moore, P. D. (1993) Btogeogruphy: A n
Ecologrcul and Evol/rtionary Approach, 5th edn, Oxford:
Blackwell.
Cronk, Q. C. B. andFuller,J. L. (1995) Pkant
Invudevs: The Threat t o Natwak Erosystems, London:
Chapman & Hall.
Darlington,
JrJ.,P. (1957) Zoogeography: The
Geographical Dtstribrrtion of Anintal.~,New York: John
Wiley & Sons.
I
Elton, C. S. (1958) The Ecology o f Inzmions by Anrnrah
und Pkunts, London: Chapman & Hall.
Hallam, A. (1995) A n Outline of Phanerozoir
Btogeography, Oxford: Oxford University Press.
Nelson, G. and Rosen, D. E. (eds) Vicariance
Btogeogruphy: A Critique (Symposium of the
Systematics Dlscussion Group of the Amerrcan
Museum of Natural History, 2-4 May 1979), New
York: Columbia University Press.
Udvardy, M. D. F. (1969) Dynamic Zoogeography'.with
Spectak Refrence to Land Anitnals, New York: Van
Nostrand Reinhold.
Wallace, A. R.( 1 876) The Geogruphiczk Distribution of
Aninmks: uuth A Study o f the Rekations o f Living and
Extinct Faunas as Ekunduting the PastChunges of the
Earth's Suvface, 2 vols, London: Macmillan.
Woods, C. A. (ed.) (1989) Btogeogruphy of the West
Indies: Past, Present. and Future, Gainesville, Florlda:
Sandhill Crane Press.
 4

POPULATIONS

Most species exist as grot+ of interbreeding individrrals - populations. This chapter covers:

the form and function of single populations

ways i n w h i c h p o p u l a t i o n s survive
human exploitation and control of populations

B I R T H ,S E X ,A N DD E A T H : THE covers the only way of ‘leaving the world’ - dylng.

D E M O G R A P H Y OF S I N G L E Death may occur through old age (senescence). It is
POPULATIONS more likely to result from disease, starvation, or
predation.Two aspects of reproduction are distin-
Populations guished. Fertility measures the actual numberof new
arrivals in a population. In humans, the fertility rate
Organisms are all to someextent sociable beings,
averages about one birth every eight years per female
interacting with one another to survive. Their inter-
ofchildbearing age. Fecundity measures the potentlal
actions create populations and communities. Popu-
number of new arrivals. In humans, the fecundityrate
lations are loose collections of individuals belongmg
is about one birthper nine toeleven months per female
the same species. Red deer (Cervrrs elaphas) In Britain
of childbearing age. Not many women fulfil that
constituteapopulation. All of themcouldInter-
particular potential!
breed, should the opportunity arise. In practice, most
Most classic population studies focused on natality
populations,includingthe red deer population in
andmortality,andassumedthatimrnrgrationand
Britain, exlst as sets of local populations or demes.
emigration balance another or are too smallto
A local red deer population lives in the grounds of contribute greatly to population change. Obviously,
Lyme Park,Cheshire.Itsmembers form atightly
emigration and immigration are important compo-
linked,interbreedinggroupand display
features
nents of long-term biogeographical change.They
typical of many populations (Box 4.1). are also important in more recent population studies
thatemphasize small-scale commgs and
goings
Population growth within aspecies range. This idea, in the guiseof meta-
population theory, will be discussed later.
A population changes due to natality and immigra-
tion,mortalityandemigration.Natality covers a
Exponential growth
variety of ways of ‘coming into the world’ - live birth,
hatching,germinating,and fission. Inmammals, Populations grow exponentially when the population
natality is the same thing as the birth rate. Mortality increase per unlt time is proportional to population
 POPULATIONS 85

Box 4.1

THEREDDEER POPULATION IN apparentgestationperiod(tothemediandate ol

LYME PARK, CHESHIRE, ENGLAND calving) was about 237 days. Stags from the Knott
rutted on Cluse Hay, while those from Cage Hill
LymeParkliesonthewestern flanks of the preferred the moor or the park. Moor stagsappeared
Pennines.Its 535 haconsist of parkgrassland, to play but a small part in the rut, despite theif
moorland, and woodland. Red deer (Cervw eluphas) numerical abundance. Dominant stags, which com-
have lived in Lyme Park for over 400 years. They are monly had 40-hind harems for two to three week:
a remnant of a larger population once present in during the rut, were usually replaced within twc
MacclesfieldForest, whichusedtoextendmany years.
kilometres along the west side of the Pennines. U p
to 1946, while the park was privately owned, the
population numbered 170 to 242. Some deer were
shot for sport and some for the dinner table. After
ownership waspassed to the National Trust and
Stockport Metropolitan Borough Council, the pop-
ulation rose to over 500. Culling was introduced in
1975 and the population is now maintained at 250
to 300.
A study of the red deer population was carried
out from 1975 to 1983 (Goldspink 1987). The pop-
ulation was dominated by hinds. There were over
threetimesmorehindsthanstags.Hinds were
recorded in three main areas - the park, the moor-
land, and Cluse Hay (which is a subdivision of the
moorland) (Figure 4.1). All tagged hinds remained
close to theirareas of capture. Hinds tended to form
large groups of 30 or more throughout the study
two weeks of
area. Most calves were born in the first
June. Maximum densities of hinds and calves per
l"2
KIII
____
:"l z n :- -I" "-1-
varlru lrum >u 111 L I K parlr --
LU
77 -- -I"
I I UII LIK
----
muur-
land. Three bachelor herds of stagswerepresent.
Moor stags spent much of the summer on a small ~

area (30 ha) of ground above Cluse Hay. They dis-

persed more during
winter,
probably avoid 4.7 The main occupjed by the red deer
sureandwindchill.TheKnottgroup,which lives in (cerous,,ldpba,) population, L~~~ park, Cheshire,
the park;consisted of primestags,morethan 6 England.
years old, during the winter. Younger stags were s o m e : After Goldspink (1987)
recorded on Cage Hill. Stags in the park area were
more tolerant of disturbancethanthoselivingonGrowthrates of stagsandhindswerelowonthe
themoorland.TherutstartedinmidSeptembermoorland. Calf-to-hindratiosweregenerally low
and went on well into November. The median date and within the range 0.20-0.34; maxima of 0.55-
of observedcopulations was 16 October.The 0.60 were recordedin woodlandsites.Conditions on
86 POPULATIONS

he moorland were severe during the winter due to
he poor quality of herbage, mainly purple moor-
p s s (Molinia cuerrdeu), and a lackof shelter. Annual
:alf mortality varied from 10 to 30 per cent. It was
)articularly high during the cold winters of 1978-9
md 1981-2. Populationdensityandwinter food
were probably principal factors limiting the perfor-
L
1
mance of deer on the moor. Growth rates of stags
and hinds in the park were high, but calf-to-hind
ratios were low at 0.15-0.26. Culling has reduced
levels of natural mortality but further improvements
in performance are unlikely to be achieved without
a reduction in sheep stocks, some improvement in
habitat, and the provisionof shelter.

size and is unconstrained. This IS readily appreciated lation (R. N. Chapman 1928). It is described by the
by a simple example. Take two individuals. After a solution to the population growth equation:
year they have produced four offspring. Reproducing
at the same rate, those four offspring will themselves N = N,e"
have produced eight offspring after a further year. The
growth of the populatlon follows a geometric progres- Thls equation shows that the startlng populatlon, No,
sion - 2 , 4 , 8 , 1 6 , 3 2 , 6 4 , and so on. This exponential increases exponentially (e is the base of the natural
growthrapidlyproduceshugenumbers.Afemale logarithms) at rate determined by the intrinsic rate
common housefly (Muscadomestrca) lays about 120 of natural increase, r. It describes a J-shaped curve
eggsatatime (LelandOssian Howard,citedin (Figure4.2).Themaximumintrinsic rate of
Kormondy 1996: 194). Around half of these develop natural increase varies enormously between specles.
into females,each of whlch potentlally lays 120 eggs. In units of per capita per day, I t is about 60 for the
The number of offspring in the second generation bacterium Escherzchzu coli; 1.24 for the protozoan
would therefore be 7,200. There seven are generations Paramecium aurelia; 0.015 for the Norway rat (Rattus
per year. If allindividuals of the reproducing genera- norveginrs); 0.009 for the domestlc
dog (Cams
tion should die off after produclng, the first fertile domestrcus);and 0.0003 for humans. For the red deer
femalewillbe a great,great,great,great,great population In Lyme Park, from 1969 to 1975, I t was
grandmother to over 5.5 trillion flies. 0.07696.
Growth in populations with overlapping genera- The doubling time is the tlme takenfor a popula-
tions and prolonged or continuous breeding seasons tion to double its w e . I t maybecalculatedfrom
may be described by the equation: values of r. The doubling tlmes for the species men-
tioned above, assuming geometric increase prevails,
dN/dt = rN are: Escherichia coli - 0.01 days; Paramiurn aurelia -
0.56 days; Rattus norveginrs - 4 6 days; Canrs domesticus
Notice that the growth rate, dNhft, is directly propor- - 77 days; and humans- 6.3 years. For LymePark red
tional to population size,N , and to the intrinsic rate deer, it is 9.0 years.
of increase (per capita rate of population growth), r.
The intrinsic rate of increase, sometimes called the
Logistic growth
Malthusian parameter, isdefined as the specific
birth rate, b, minus the specific death rate, d. In sym- The Englishhousesparrow (Passwdomestrcus) was
bols, r = b - d. (Specific rates are rates per individual introduced to the United States around 1899.Concern
per unit time.)Exponentialgrowth,then,slmply was expressedthat in a decade, a slngle pair lead could
dependsuponhowmanymorebirthsthereare to275,716,983,698descendants,and by 1916to
than deaths, and on how large the population is. N o 1920therewould be about575birdsper 40 ha
other factors restrict population growth. Unhindered (Kormondy 1996: 194). In the event, there were only
growth of this kind IS the biotic potential of a popu- 18 to 26 birds per 40 ha, less than 5 per cent of the
 :
POPULATIONS 87

Exponential population growth logistic population growth

I N . K

I
N = N&" 1 + ce"'
Carrymg capaclty. K I 3.000
Starting populatlon. No = 2
Slartlng populatlon. No = 2
Constant. C = ( K / N o ) - l = 1.499
Growth rate. r I 0.3

/ 1
3.000
+ t.499-03'

10 20 30 40 50 1 I1 40 50
Time, I (years)

Figure 4.2 Exponential growth of a hypothetical popula- F i g w e 4.3 Loglstic growth of a hypothetical populatlon.
tion. Notice that, wlth a bit of artistic licence, the Notice that the population growth curve is a stretched
popularlon growth describes a J-shaped curve S-shape

expected denslty. Patently, something had prevented approachingthecarrying capaclty, environmental

the unfettered geometrlc Increase in the immigrant resistance is great. The populatlon barely grows. Thls
house sparrow populatlon. 15 the distorted top part of the 'S'. If the populatlon

A growing populatlon cannot increase Indefinitely
at a geometric rate. There is a ceiling or carrying
capacity, a limlt to the number of individuals that a
gwen habitat can support. The biotic potential of a
populatlon I S curtailed by an environmental rem-
tance (R. N. Chapman 1928). After a period of rapid
growth,the housesparrow population reached its
population ceiling, partly because native hawks and
owls took to the new English dish on the menu.
Thepopulationceiling or carrying capacity is
usually denoted by the letter K. It alters the popula-
tlon growth equation in this way:
JNldt = rN( I - NIK)
This equation for growth rate is called the logistic
equation. It describes a sigmoid or S-shaped curve
(with the top of the 'S' being pushed to the right)
where population grows fast initially but tapers off
towards the carrymgcapacity (Figure 4.3). When the
population size I S small, compared wlth the carrylng
capacity, the environmental resistance to growth is
minimal.Thepopulatlon responds by virtually
unbounded exponentlal growth. This IS the bottom
part of the 'S'. When the populatlon S I X is large and
should exceed the carrying capacity, negative growth
occurs;in other words, the populatlon falls until I t
drops below the carrylng capacity.
Populationgrowth (or decline) rates are affected
by density-dependentfactorsand by density-
independent factors.
The effects of density-
Independent factors donot vary wlthpopulation
density and theywill affect the same proportion of
organisms at any density. Weather, pests, pathogens,
andhumanscommonly affect populations in that
way. The effects of density-Independent factors do
vary withpopulationdensity, so thattheportion
of organisms affected increases or decreases with
population density. Birth rates and death rates both
dependonpopulatlondensity:birth rates tendto
decrease wlth increasing density,whiledeath rates
tend to Increase with Increasing density.
Population irruptions
Irruptions are common features of many populatlons.
They Involve a population surgmg to a hlgh density
andthen sharply declining.Thisboom-and-bust
pattern occurs in some introduced ungulates. It hap-
pened to the Himalayan tahr (Hrrtlitraps jemlah/cx.r)in
88 POPULATIONS
New Zealand (Caughley1970). The tahris a goat-like
ungulate from Asia. Introduced into New Zealand In
1904, it spread through a large part of the Southern
Alps. After its Introduction, the population steadily
rose. The birth rate fell slightly but the death rate
increased,largely due to greater juvenile mortality.
After a perlod when the populatlonwas large, a decline
set In. This declinewas caused by reduced adult fecun-
dity and further juvenile losses. Changes In the tahr
populatlon may have resulted from changesin food
supply. As the tahr grazed, they altered thecharacter
of thevegetationthatthey fed on. In particular,
snow tussocks (Cbtonoc-hlou spp.), whlch are evergreen
perennial grasses, were an important food source late
In winter. They cannot abide even moderate grazing
pressure and disappeared from land occupied by the
tahr. With no snow tussocks to eat, the tahr browsed
on shrubs in winterandmanagedtokillsome of
these.
Another
Irruption occurred
on Southampton
Island, Northwest Terrltories, Canada (D. C. Heard and
Ouellet 1994). By 1953, Caribou (Rangifrv tavandw
groenlandicrrs) had been huntedtoextinctlon on
Southampton Island. In 1967, 48 caribou were
captured on neighbouring Coat’s Island and released
on Southampton Island. The population of caribou
older than 1 year grew from 38 In 1967 to 13,700 in
1991. The annual growth rate during this period was
27.6 percent and did not decrease wlth increasing
population density. On Southampton Island, caribou
did not suffer high wlnter mortalityIn some years, but
they did onCoat’s Island. Caribou density was higher
on Coat’s, which suggests that adverse weather has a
mlnlmal effect when animal density is low.
Irruptions also occur because of unusually high
immigration. In the winter of 1986-7,manymore
rough-legged buzzards ( B ~ t e o/app//.r) moved Into
Baden-Wiirttemberg, south-west Germany, than in
any other winter durlng atleast the preceding century
(Dobler et a/. 1991). From 14Januaryto7April,
rough-legged buzzards were observed daily.The
highestnumbers observed were 109 indivlduals on
1 February and 110 on 8 February. The buzzard Influx
was caused by high snow cover and cold spells In the
eastern parts of CentralEurope. Depresslons over
Southern Europe may have blocked the route south
wlth cloud and snowfall.
Population crashes
.%me populations often crash. In early 1989, two-
thirds of the Soay sheep (Oz~isa r k ) population on
StKilda, In theOuterHebrides,Scotland,died
within a twelve-week period. The cause of the crash
was investlgated by post-mortemexamlnationand
laboratory experiments (Gulland 1992). Post-mortem
examinatlon showed emaciated carcasses with a large
number of nematode (Ostevtagta Lwum/cincta) parasltes,
andthattheprobable cause of death was proteln-
energy malnutritlon. However, well-nourlshed Soay
sheep artificiallyinfected withthe paraslte in the
laboratory showed no clinlcal slgns o r mortality, even
when their paraslre burdens were the same as those
recorded in the dead St Kilda sheep. Thus, parasites
probably contributed to
mortality only in mal-
nourishedhosts, exacerbatingthe effects of food
shortage.
Marine iguanas (Amhlyrbyncbrrs c.rtstaut/rs) living on
Sante Fe, in the GalipagosIslands, suffered a 60 to 70
per cent mortalityduetostarvationduringthe
1982-3 El Nitic-Southern Oscillation event (Laurie
and Brown 1990a, b). Adult males suffered a hlgher
mortalitythanadult females while food was short,
but size explained most of the mortality differences
between the sexes. Almost no females bred after the
event. In the next year, the frequency of reproductlon
doubled, theage of first breeding decreased, and mean
clutch s ~ z eIncreased from two to three. These demo-
graphic changes are examples of denslty-dependent
adjustments of populationparametersthathelpto
compensate for the crash.
Population crashes may have repercussions withln
communltles.In Afrrlca, savannahsare malntalned
as grassland partly because of browslng pressures by
large andmedium herbivores. In Lake Manyara
Natlonal Park, northern Tanzania, bushencroached
on thegrassland between 1985 and 1991, shrubcover
Increasing by around 2 0 per cent (Prins and Van der
Jeugd 1993). Since 1987, poaching has caused a steep
decline in the Atrlcan elephant (Loxodonta afrtcum)
 POPULATIONS 89

population in thePark. However, shrubestablish- and Wedin 1991). Monocultures of tlcklegrass were
ment preceded the elephant population decline, and planted at two different densities on ten different soil
I S not attributable to a reduction in browsing. In two mixtures. Populations growing in unproductive soils,
areas of thePark,shrubestablishment colncided low in nltrogen,mamtained fairly stable above-
withanthraxepidemicsthatdecimatedtheimpala ground biomass. Populations growlng on richer soil
(Arpyceros ttdattzpus) populatlon. In
the
northern showed biomass oscillations. Populations growlng on
section of the Park the epidemic was in 1984; in the the rlchest soils displayed as 6,000-fold crash (Figure
southern sectlon it was in 1977. An even-aged stand 4.5). No other species growing in the garden where
of umbrellathorn (A~aciurovti1z.r) was established the experimentwas conducted crashed in 1988,whlch
wlthin the grassland In 1961, which date coincided suggests that environmental agencies were not respon-
wlthanother
anthrax
outbreak
amongmpala. sible. Thedynamics for thecrashingpopulations
Similarly, another even-aged stand of umbrella thorn were shown to be chaotlc. The chaotlc regime resulted
was established at the end of the 188Os, probably fol- from the growth inhibltlon by litteraccumulation.
lowing a rinderpest pandemic. The evldence suggests The littercauses a one-year delay between growth and
that umbrella-thorn seedling establishment is a rare the lnhibltlon of future growth. Litter production is
event, largely owlng to high browsing pressures by greater In more productive plots. The magnitude of
such ungulates as impala.Punctuateddisturbances the time-delayed inhibltory effect therefore increases
by epidemlcs, which cause populatlon crashes In withproductlvity.Thls may lead to anoscillatlon
ungulatepopulations, createnarrow windows for between a year of low litter and hlgh living biomass,
seedling establishment. This process may explaln the and a year of high litter and low living blomass. The
occurrence of even-aged umbrella thorn stands. lmplications of thls study are profound - litter feed-
back may cause population oscillations and possibly
chaos in productlve habitats. However, litter accumu-
Chaotic growth
lates where I t falls, and litter-driven chaotic dynamlcs
Irregular fluctuatlonsobserved in natural populatlons, occurs at small spatlal scales - it may avoid detectlon
including explosions and crashes, were traditionally if looked for at medium and large scales.
attributedto external random Influences such as Since the late 1980s, partof theoretical ecology has
climateand disease. Inthe 197Os, RobertM. May focused on thespatio-temporaldynamicsgenerated
recognized that thesewild fluctuations couldresult by simple ecological models. To a large extent, the
from intrlnsic non-linearities in populatlon dynamics results obtained have changed views of complexity
(May 1976). For instance, a relatively slmple populatlon (Bascompte and Sole 1995). Specrfically, slmple rules
growth equatlon of the form are able to produce complex spatlo-temporal patterns.
Consequently,some of thecomplexltyunderlylng
,
N,, = N,~[I'(1 - N,/K)] Nature does not necessarily have complex causes. The
emerglng framework has far-reaching implications
which I S applicabletopopulatlonthat suffer epi- in ecology and evolutlon. It I S Improvmg the compre-
demlcs at hlgh densities, possesses an amazing range henslon of such t o p m as the scale problem,the
of dynamlc behavlour - stable points, stable cycles, response to habltatfragmentation, the
relatlon-
and chaos - depending on the growth rate, I' (Figure ships between chaos and extinction, and how higher
4.4). diverslty levels are supported in Nature.
Much of the work on chaoticpopulation
dynamics I S theoretlral. Some studies have shown
Age and sex structure
that chaos does occur In naturalpopulatlons.The
tlcklegrass or hairgrass (Ayostis sm!wu) displays Theaggregatepopulatlon slze, N, obscures the
oscillatlons and chaos In experimental plots (Tilman fact that Indivduals differ In ageand in ability to
90 POPULATIONS

-
(a) Stable equilibrium point Growth rate, r = 1.8

1-

0
2 1 (b)Stable two-point cycle Growth rate, r = 2.3

0"
i l
2 -- (c)Stable four-point cycle Growth rate, r = 2.6

0.
S 31 ( d )Chaos Growth rate, r = 3.3

-
3 1 (e) Chaos Growth rate, r = 3.3

12 7 ( f ) Chaos Growth rate, r = 5.0

1
9-

1 1 1
0 10 20 30
Time, f

Figure 4.4 Inrernally driven population dynamics: stable points, stable cycles, and chaos. NB The growth rates, r, in (d)
and (e) are the same, bur the rarios of rhe Initial population to the carrying capacity, N,lk, are different: i n (d), N[/k =
0.075; In (e), N,lk = 1.5.
Sorrrce: After May (1981)

reproduce. The populationage-structure is thepro- Life tables

portion of individuals in each age class. A stable age
distributlon with azero growthrate has about the
same number of individuals in each age class, except
for fewer In post-reproductive classes. In contrast, a
growing population has a pyramid-shaped structure,
with large numbers of pre-reproductive indivlduals.
A life table, like an actuarial table, shows the num-
ber of individuals expectedin each age classbased
uponfecundityandsurvival rates. Survival rates, I,
arerelated to death rates, d, inthefollowlng way:
I = 1 - d. Life-table sratistlcs are normally computed
 POPULATIONS 91

800 I 1,200 T 0 N1
V N2
A N3
N
E N4
- 600
Ln

m 800
.-E
I)

7 400
3

& al 400
0
n
< 200

0 0
< ds
1990
1989
1988
1987
1986

4 . 5 A population crash In the r~cklegrass(Agrmtrs .r(.uhru).(a) High seed density monocultures. (b) Low seed denslty
F;rpr~e
monoculrures. The soil mlxrures are divided into four g r o ~ ~ pNl-N4.
~~
s, Group N1 is poorest in nitrogen, and Group N4
IS the richest In nirrogen.
Sorrrce: After Tilman and Wedin (1771)

for females. This is because fecundity for males I S
very difficult todetermine where mating occurs
promiscuously. A life table for a hypothetical popu-
lation of females IS shown in Table 4.1. Column 1
lists the age classes. Column 2 gives the survivorship
expressed as the fraction survivlng at agex; this is the
probability that an average new-born will survive to
that age. Column 3 gives the fecundity; this is the
number of offspring produced by an average individ-
ual of age x during that period. The sumof fecundity
terms over all ages, or the total number of offspring
that would be produced by an average organism with
no mortality, is called the gross reproductive rate. It
is 9.8 in the example. Column 4 shows the number
of expectedoffspring by age class. The figures are
simplytheproduct of survivorshipandfecundity.
The sum of expected offspring gives the net repro-
ductive rate, R. Thls is the expected number of
femaleoffspring to which a new-bornfemalewill
give birth in her lifetlme. I t is 7 . 3 2 2 in the example.
Column 5 shows the product of age and expected off-
spring. The totalof these products divided by the net
reproductwe rate defines the mean generatlon time,
T . This is the average age at which females produce
offspring. I t is 5.26714 years In theexample.The
earlier young are born, the earlier they, in turn, will
have offspring and the more rapid population growth
willbe. The calculations show that the population
will increase by a factor of 7.322 times every 5.26714
years. Plainly, growth rates expressed on a generation
basis would be difficult tocompare. To avoid this
problem, growth rates are normally converted to an
annual figure anddesignated by theGreekletter
lambda (1). Theannualgrowth rate of the hypo-
thetical population is 1.459.
Survivorship curves
A survivorship curve shows the fraction of a cohort
of new-born (or newly hatched) alive individuals in
subsequent years. Naturalpopulations have a greac
range of survlvorship curves. Three basic types are
recognlzed (Pearl 1928) (Figure 4.6):
1 Type I survivorship curves are 'rectangular' or
convex on seml-logarithmic plots. They show low
92 POPULATIONS

mortality Initially that lasts for more than half the
life-span,after whlchtimemortality increases
steeply.Thissurvivorshlppattern I S common
amongst mammals, including the Dall mountain
sheep (Oz~icdalli dalfi) and humans. It is also dis-
played by such reptiles as the desert night lizard
(Xantrtsia vrgih).
2 Type I1 survivorshipcurves are ‘diagonal’ or
straight on semi-logarithmic plots. They show a
reasonably constant mortality withage.
Thls
survlvorshippattern is common inmost birds,
Including the Amerlcan robin (TIodus rnrgratorrus
rnrgratorlzu), and reptiles.
3 Type 111 survivorshipcurves are concave on
semi-logarithmic plots. Theyshow extremely high
juvenile moralityand relatively low mortality
thereafter. This survivorship pattern I S common In
many fish, marine Invertebrates, most Insects, and
plants. I t is also characteristlc of the British robln
(Errtbaus rrrberxfa nzefopbifuc).
These three basic survlvorship curves do not exhaust
all the possibilities - many intermediate types occur.
Cohort-survival models
Models of population disaggregated by age and sex
arecalled cohort-survivalmodels (Leslie 1045,
1948). A cohort-survivalmodel starts wlth the age
structure of a female population at a given time, and
with the blrth rates and survival rates per lndivldual
In each age group. From thls informatlon it predicts
the age structure of the female population in future
years.
Cohort-survival models are used to study popula-
tion change. One applicatlonconsidered the effects of

Table 4. I Life table of a hypothetical population

Column 1 Column 2 Column 3 Column 4 Column 5

Age,
(years)
x Survivorship, /x Fecundity, bx Expected
Product of age
offspring, lxbx and expected
offspring, xlxbx

0 1 0 0 0
1 0.99 0.2 0.2 0.2
2 0.98 0.4 0.392 0.784
3 0.96 0.8 0.768 2.304
4 0.92 1.2 1.104 4.4 16
5 0.86 1.6 1.376 6.880
6 0.78 2.2 1.716 10.296
7 0.68 1.4 0.952 6.664
8 0.56 0.8 0.448 3.584
9 0.42 0.6 0.252 2.268
10 0.26 0.4 0.104 1.040
11 0.06 0.2 0.012 0.132
12 0 0 0 0
9.8 7.322 38.546
This is the gross This i s the net This is the
reproductive reproductive total weighted
rate, GRR rate, R age

Mean generation time, T = 38.566/7.322 = 5.26714 years

Annual growth rate, A, is given by h = R’IT. It is 7.322’/5.267’4
= 1.459
 POPULATIONS 93

( a) 1 ---
1,000

800
Type I curve
Dall mountam sheep

g 600

.
L
0

n
f 400
z

200 \ ... \ 1 \.'

0
6020 40 0 BO 100 100 80 6020 40
Per cent of total
Perlifespan cent of total lifespan

over-exploltatlonon thebluewhale (Balurnopteru of growing. The IntrlnsLc rate of natural Increase of

m/m./////s)population(Usher1972). It used dataon
thebluewhalepopulationcollected in the1930s,
before the near extlnction of the species In the early
1940s. Agespecific birth rates and survival rates for
each of seven age groups were calculated (Table 4.2).
The birth terms (fecundity terms in this study) show
that female blue whales clo not breed in the first four
years of life. At full breeding, each cow produces one
calf every two years (since the sex ratlo I S 1:1, thls
calf has a 50 per cent chance of belng female). The
survival terms, all but one of which is 0.87, I S based
on the bestavailableestlmateofnaturalmortality.
The survival rate of the 12-plusage class IS 0.8. Thls
is because not all members of the oldest age-group
die wlthin a two-year period; indeed, some live to be
40. The survival rate of 0.8 per two years gives a life
expectancy of 7.9 years for old whales.
The parameters in the table may be arranged as a
growth matrlx,
from
whlchthe
growthcharac-
teristics of the whale populatlon may be extracted.
The dominant latent root, h,, of the growth matrlx
IS 1.0986. This shows that the population is capable
thebluewhalepopulation is given by thenatural
logarithm of the dominant latent root;I t is In 1.0986
= 0.094. The dominant latent root may be used to
estimatethe harvest of whales that can betaken
withoutcausingadeclineinthepopulation.The
populatlon slze increases from N to h , N over a two-
year period.The harvest that may betaken, H,
expressed as percentage of the total populatlon, I S
H = 100 [(h,-l)/h,]
This is about 4.5 per cent of the total population per
year. If the harvest rate were exceeded, the popula-
tlon would decrease, unless homeostatlc mechanisms
should come into play and alter fecundity and sur-
vival
rates.
Evldence
suggests that,whenunder
pressure, thebluewhalepopulatlonshowsaslight
increase inthepregnancyrateandthatindividuals
reach maturltyearlierin life, andthatthey may
generally grow more rapldly.The big questlonIS: can
these responses offset the effects of exploitation? For
a trustworthyanswer to thlsquestlon,long-term
94 POPULATIONS

Table 4.2 Parameters for a cohort-survival model of the blue whale (Balaenoptera musculus)
population

Parameters Age groups

(per cow per
2 years) 0- 1 2-3 4-5 6-7 8-9 10-1 I 12+

Birth rate 0 0 0.1 9 0.44 0.50 0.50 0.45

rate
Survival 0.87
0.87
0.87 0.87 0.87 0.87 0.80
Source: After Usher (1 972)

studies involving data collection and modelling are
11eeded.
Thewanderingalbatross (Dlotnediu exlrluns) is a
splendid bird, with a wing span of over three metres.
It breeds on southerncool-temperateandsub-
Antarctic islands, and forages exclusively in the
Southern Hemisphere. Wanderlng albatross popula-
tions around the world have declined since the mid-
1960s. The major factor contributing to this decline
is accldental deaths associated with longline fishing,
which has increased rapidly since the 1960s. Entan-
glementand collisions withnetsondemonltor
cables ontrawlers may also contribute to albatross
mortality. Longline operations involve baited hooks
attached to weighted lines being thrown overboard.
Seabirds lured to the bait swallow the hooks and die
by drowning when they are dragged beneath the sur-
face as the weighted line sinks, or die when the lines
are hauled. Some 44,000 albatrosses, including about
10,000 wandering albatrosses, are killed in this
manner each year (Brothers 1991). An age-structured
model of a wandering albatross population, based on
demographic data from the population at Possession
Island, Crozets, simulatedpopulationtrends over
time(Mo1oney etal. 1994).Theaim was to investigate
the potential Impacts of new longline fisheries, such
as that for thePatagonian toothfish (Dissosticbus
eleginoides) in Antarctica. Themodel consisted oftwelve
year age-classes, comprising four age categories:
chicks (prior to fledging, 0-1 years), juveniles (before
the first returntothebreeding colony, 1 4 years),
immatures (after returning to the colony but before
the first breeding, 5-10 years), and breeding adults
(more than 10 years). The model kept track of the
population in each age class, N,.Thepopulations
change owing to eggs being laid (fecundity) and sur-
vival from one age class to the next. Fecundity and
survival terms are summarized In Table 4.3, together
with starting numbers for a stable population. The
simulationspredicteda decreasing population. The
rate of decrease is constantat 2.29 per cent per
year and is assoclated withastableagestructure
of 14 per cent chicks, 24 per cent juveniles, 15 per
centimmatures,and 47 per centadults.Further
simulations werecarrled outtotestthesensitivity
of the wandering albatross population to altered sur-
vival rates. On the assumption that longline fishing
operations affect juveniles more than adults, thereis a
time lag of five to ten years before further decreases in
population numbers are affected in the breeding pop-
ulations. In addition, because wandering albatrosses
arelong-lived,populationgrowth rates takeabout
thirty to fifty years to stabilize after a perturbation,
such as the introduction of a new longline fishery.
Metapopulations
There arefourchief types of metapopulations,
normallystyledbroadlydefined,narrowlydefined,
extinction and colonization, and mainland and island.
The broad and narrow categories are more concisely
named loose and tight metapopulations, respectively.
Loose metapopulations
A loose metapopulation is slmplya set of sub-
populations of thesame species. Rates of mating,
competition, and other interactlons are much higher
 POPULATIONS 95

Table 4.3 Demographic parameters for a wandering albatross (Diomedia exulans) population

Age class Age category Survival Starting population

(Years] (per Year] (for a stable age
structure]
~
~ ~~

0 Eggs and chicks 0.640 313

1 Juveniles 0.71 5 20 1
2 Juveniles 0.715 147
3 Juveniles 0.715 107
4 Juveniles 0.715 78
5 lmmatures 0.980 57
6 lmmatures 0.980 58
7 lmmatures 0.980 58
8 lmmatures 0.980 59
9 lmmatures 0.980 59
10 lmmatures 0.980 60
11 Adults 0.922 1,066

Source: After Moloney et a/.( 1 994)

wlthlnthesubpopulatlonsthan theyarebetween populations stabilizes local population fluctuations.

thesubpopulatlons.Thesubpopulatlons may a r w This 'rescue effect' helps to prevent local extlnctions,
through anaccldent of geography - thehabitat throughthe recolonizatlon of new habirats or
patches in which the subpopulations live lie farther habitats lefr vacant by local extinctlons (or both). In
apart than thenormal dispersal distance of the species. theory, a tight metapopulatlon structure occurs
The conspecific subpopulationsthatcomprise such where the distance between habitat patches is shorter
a metapopulation may or may not be interconnected. than the species is physicallycapable of travelling,
By thisdefinition, most species with large and but longer than the distance most individuals move
discontinuous distributlons form metapopulations. within their lifetimes. An example is the European
Where habitat patchesare so close together that nuthatch (Sitta europaea) population.Thenuthatch
most individuals visit many patches in their lifetime, inhabitsmaturedeciduousand mixed forest and
the subpopulations behave as a single population - has a fragmenteddistribution in theagricultural
all individuals effectwely live together and interact. landscapes of western Europe (Verboom et al. 1991).
Where habitat patches are so scattered that dispersal It showsall the characteristics of a tightmeta-
between them almost never happens, the subpopula- population:thedistribution I S dynamicin space
tions behave effectlvely as separate populations. This and time; the extlnction rate depends on patch size
situationappliestomammalpopulationsliving on and habitat quality; and the patch colonization rate
desertmountain-tops in theAmericanSouthwest depends on the density of surrounding patches
(Brown 107 1). occupied by nuthatches.

Tight metapopulations Extinction-and-colonization metapopulations

A tight metapopulation is a set of conspeclfic S U ~ - In the narrowestpossible sense, and as orig~nally
po~x~lationsliving In a mosaic of habitat patches, defined (Levins 1970), a metapopulation consists of
with a slgnlficantexchange of individuals between subpopulations characterized by frequent turnover. In
the patches. Migration o r dispersal among the sub- these 'extinction-and-colonization' metapopulations
96 POPULATIONS
Figure 4.7 T h e Granville fritillary (Melituu anxru) populatlon In &and, Finland,
meadows; solid clrcles are occupled meadows.
Sourre: After Hanskl et ul. (1996b)
(GutierrezandHarmon 1996), allsubpopulatlons
areequallysusceptibletolocalextmction.Specles
perslstence therefore depends on there bemg enough
subpopulatlons,habitatpatches,anddispersalto
guaranteeanadequaterate of recolonization.This
metapopulation structure seems to apply to the pool
frog (Kana fmonae) on the Balm coast of east-central
Sweden (Sjogren 1991).Frog lmmrgratlon mmgates
agarnst large population fluctuatlons and inbreeding,
and 'rescues'local populatlons
from
extrnctlon.
Extmction-and-colonizatlon metapopulatlon structure
also applies to an endangered butterfly, the Glanville
in 1993.Shaded clrcles are empty
frrtillary (Melitaea crnxza) in a fragmented landscape
(Hanskl et a / . 1995; 1996b). Thls butterfly became
extinct on the Finnish mainland in the late 1970s. It
is now restrlcted t o t h e k a nIslands.
d It has two larval
host plants on the Islands - ribwort plantain(P/antago
fanceolata) andspikedspeedwell (Veronicaspzcata)
- both of which grow on dry meadows. The meta-
populatlon was studied ona network of 1,502 discrete
habitat patches (dry meadows), comprlsing the entwe
distributlon of this butterfly specles In Finland (Figure
4.7).Survey of the easilydetectedlarvalgroups
revealed a local populatlonIn 536 patches. The butter-

fly metapopulation satistied the necessary conditions
for a species to persist in a balance between stochastlc
local extinctions and re-colonizations.
Mainland-island metapopulations
In some cases, there may be a mixture of small sub-
populations prone to extinctlon anda large persistent
population. The viability of the ‘mainland-island’
rnetapopulations (GutiCrrez and Harrlson 1996) is
less sensltlve to landscape structurethan In other
types of metapopulatlon. An example IS the natter-
jack toad (B& l-alunlrtu) metapopulationat four
neighbouring breeding sites In the North Rhineland,
Germany (Sinsch 1992).Over 90 per cent of all
reproductlve males showed a lifelong fidelity to the
site of first breeding; females did not prefer certain
breeding sites. Owing to the female-biased exchange
of individuals among neighbouring sites, the genetic
distance between local populations was generally
low but increased withgeographicaldistance.This
spatlal pattern is consistent with the structure of a
mainland-island metapopulation. Up to three mass
immigrations of males per breeding period, replacing
previously reproductiveindividuals,suggestedthe
exlstence of temporal populations successlvely repro-
duclng at the same locality. Genetlc distanceswere
considerably greater between temporalpopulations
than between local ones, indicatingpartlal repro-
ductive
isolation.
An exchange of reproductlve
Individuals between thetemporalpopulationsat
each site was notdetected,butgene flow dueto
therecruitment of first-breeders ortglnating from
offspring otherthantheirown seemed probable.
Thus,natterjackmetapopulatlons consist of m e r -
actlng local and temporal populations. Three out of
four local populations had low reproductive success,
as well as the latest temporal populatlon. The persis-
tence of these populations depended entirely on the
recruitment of juveniles from the only self-sustaining
local population.Thls ‘rescue-effect’ impeded local
extinction.
POPULATIONS 97
Metapopulations and conservation
MetapopuIat1ontheory is now wldely applied in
conservation brology. I t I S especially important where
habitat fragmentation is thecentralconcern.The
‘classic’ example is the northern spotted owl in the
UnitedStates, for whichbird thcmetapopulation
approach to management has been thoroughly devel-
oped.Other examples abound,includingtheeuro
and Leadbeater’s possum, both from Australia.
The spotted owl in north-western North
America
The northern spotted owl (Strm ocudtntalis caurit/a) is
closely assoclated with mature and old-growth con-
iferous forests In the Pacific Northwest. These forests
onceformed acontinuoushabltat,but they have
shrunk over the past half century. They now survive
as small remnants occupying ~ u s t20 per cent of their
former extent. This fragmentation of the spotted owl
habitat reduces dispersal success, whlch may jeopar-
dize the long-term survlval of the spec~es. Northern
spotted owls require terrltorles of around 1,000 ha.
They avoid open areas, where they are vulnerable to
predatlon by the larger great-horned owl (BNIIo
zirginianu.r), and will cross only small cleared strips.
Thelr survival therefore depends on the connectwity
of the forest habltat (Ripple et dl. 1991). The preser-
vatlon of a viable owl population I S requlred by the
Endangered Species Act and by laws governing the
United States Forest Service, whlch owns 70 per cent
of the owl’s remalnlnghabltat.TheUnited States
government commissioned scientlsts to discover
how much forest, and in whatform, was requlred
to ensure the owl’s survlval. The result was a spatial
model of the owl population (Lamberson et a/. 1992;
see also S. Harrison 1994; GutlerrezandHarrlson
1996).
The modellet a single spatial element represent
the terrltory of a pair of owls. The dynamlcs of each
pairweremodelled usmg rates of pair-formation,
fecundity,
and survlval
measured
in the field.
Dispersal was estlmated fromjuvenileowls tagged
with radio transmitters. Owl demographywas linked
98 POPULATIONS

to forest fragmentatlon by juvenile dispersal success. lndivlduals livedalone or In small groups, movlng
To join the breeding populatlon, a newly fledged owl frequentlybetween several remnants.Overall,the
needed co find a vacantandsultableterritory. Its euros In the1,680-km'studyarmappeared to be
chances of domg so depended upon the proportion of separated Into a number of metapopulations, some of
forestedlandscape. The resultspredicted a sharp whlch had very small numbers of animals. Within
threshold,belowwhichtheowlpopulationcould themetapopulatrons,movementsbetweenpopula-
notperslst. With less than20 percentforest,the tions appear to be dependent on the availability of
juvenilerecrultment to thebreedingpopulatlon 'stepplngstones'andcorridors.Intwopopulatlons
was too low to balance mortality of adult territory that had low densitiesofeuros,thenumbers of
holders,andthepopulationcollapsed.Inaddition, juvenilesperadultfemaleweresignificantlylower
whenjuvenileswererequlred to find mates as well than insystems wlthhigherdensities.Long-term
as empty territories,a second survivalthreshold survival of these two populatlons I S questionable.
emerged, below which extinction soon ensued. The
recommendation was that 7.7 million acres (about 3
Leadbeater's possum
million ha) of old-growth forest should be preserved
in a system of patches, each patch large enough for Leadbeater'spossum (G~mnohelic/twslradhecrteri) lives
morethantwentyowlterritories wlthln12 miles In the cool, mistymountain forests of theCentral
(about 20 km) of the nearest patch Highlands, Victorla,
Australia(Colourplate
6).
These forests are timber-producrng areas. Forest frag-
mentation is threatenlng the survlval of the possum,
The euro
which is an endangered specres.
Thecommon wallaroo or euro (h1mvp.r ~ o h m t m ) , The viability of possum metapopulatmx in the
called theblggada by aborlgrnes, I S alarge,rather fragmentedold-growth forests wasassessed uslng
shaggy-hairedkangaroo(Colourplate 5 ) . Itsrange a population model(Lindenmayerand Lacy 1995).
Includes most of Australia, except the extreme south Computer simulations wlth subpopulatlonsof twenty
andthewestern s ~ d eof theCapeYorkPeninsula, or fewer possumswerecharacterized by very rapid
Queensland. Its habitat usually features steep escarp- ratesof extinction, most metapopulations typically
ments, rocky hills, or stony rlses; and, in areas where failing to persist for longer than fifty years. Increases
extreme heat is experienced for long periods, caves, In either the migratlon rate or the number of small
overhanging rocks, and ledges for shelter. subpopulations worsened the metapopulatlons' demo-
Members of a europopulationlivlng In a frag- graphic instability,at least whensubpopulations
mentedlandscapeoccupleda very large (1,196 ha) contained fewer thantwenty~ndivldualsandwhen
patchofremnantvegetatmn (G. W . Arnold et crl. migration rates were kept wlthm realistlc values for
1993). Theywere sedentary and did not use any other Leadbeater'spossumsdispersingbetweendisjunct
vegeratlonremnants.However,someyoungmales habltatpatches. The worsening demographlc sltua-
fromthisremnantdispersed up to 18 km to other tlon was reflected In lower rates of populatlon growth
remnants.In areas that had alargeremnant(more anddepressedprobabilitiesofmetapopulationper-
than 100 ha), individuals were also sedentary. Some sistence. These effects appeared to be associated wlth
individualsincluded In theirhomerangessmaller substantialImpactsofchancedemographlcevents
remnants within 700 m. Toaccess these, they usually on very small subpopulations, together with possum
moved along connectlng corridors between remnants. dispersal intoeitheremptypatches or functionally
Occaslonally,
a few indivlduals made
excursions extlnct (single-sex) subpopulatlons. Increased migration
of several kilometresoutsidethelrnormalhome rates and the addition of subpopulations containing
ranges,eitherovernight or over several months. In fortypossumsproducedhlgherratesofpopulation
areas where all remnants were small(less than 30 ha), growth, lower probabilitles of exclnctlon, and longer
 POPULATIONS 99

perslstence tlmes. Extinctlons in these scenarios were stances, there is nothing to be lost by exceeding the
also more likely to be reversed through recolonizatlon habltat's carrying capacity because the resources left
by dispersing individuals. for futuregenerationswillnot be jeopardized. For
A common problem with fragmented populatlons this reason, short-lived habitats tend to he occupied
I S thatgeneticinformatronwithinthegene pool by exploiters or opportunistswithboom-and-bust
becomes less well mlxed. Atthehlghest rates of populatlon dynamics. Such opportun1stIc species are
migratlon,subpopulatlons of fortypossumswere called r-strategists, after the lntrlnslc growth rate, r .
thoroughly mixed and behaved genetically as a slngle in population growth equatlons.
larger population. However, over a century, the gene- Another extreme case occurs where the generatlon
pool mlxing would decline and lead to a slgniticant time I S tlnycomparedwith a stablehabitat's 'life-
loss in genetic variability. This loss would occur even span'. Thiswould apply, for instance,to an orang-utan
wlthhighestmigration rates among five, forty- (Pongo p y p u e m ) , to whom an entlre forest is a stable
animal subpopulatlons. While demographlc stability and permanent habitat. Under these circumstances, it
might occurinmetapopulatlons of twohundred pays species to look to the fxure.Long-lived habitats
individuals, considerably more indivlduals than this witha falrly stablecarryingcapacitywould be
might be requlred to avoidaslgnificantdeclinein degraded if population density should overshoot the
genetlc variability over a century. carryingcapacity. Thiswouldreducethehabitat's
carryingcapaclty for futuregeneratlons. Species in
thesehabltats arc adapted to harvesting food 111 a
A Q U E S T I O N OF S U R V I V A L : crowded environment. They are hlghly competltlve
P O P U L A T I O NS T R A T E G I E S and they tend to have low population growth rates.
They are calledK-strategists, after the carryingcapac-
Anlmals and
plants possess prlmary ecological ity parameter, K , In populatlon growth equations.
strategles. These strategles are recurrent patterns of
speclalizatlon for life in particular habitats or nlches.
The r-strategists
They Involve the fundamentalactivities ofan organism,
including resource capture, growth, and reproduction. Ther-strategistscontinuallycolonizetemporary
There are two main schemes for population strategles. habltats. Their strategy I S essentlally opportunistic.
The tirst lnvolves two basic strategies - opportunism They have evolved highpopulationgrowth rates,
and competitlveness; the second involves three baslc produced by a high fecundity and a short generatlon
strategies - competitiveness,stresstoleration,and time. Migration is a major component of thelr popu-
ruderalism. lation dynamics, and may occur as often as once per
generation. The habitats they colonlze are commonly
free fromrlvals. This fiavours a lack of competitlve
Opportunists and competitors abilitlesandsmall slze. Theydefendthemselves
Two baslc populatlon strategies depend on how long against
predators by synchronizing generations
a habitat is favourable. A crucial factor is a popula- (hencesatiatingpredators)and by beingmobile,
tion's generatlon time compared with the 'life-span' which enables them to play a game of hide and seek.
of a stable habitat. One extreme case is where the gen- Thegreatestv-strategistsare insects andbacteria.
eration time is about the same as a stable habitat's The African armyworm (Spodopteru exetqbt't) is a good
'life-span'. Thiswouldapply, for example,tofruit example. This migrant moth I S a crop pestin East
flies (Drosophila)living in a tropical forest. A fruit fly Africa. It lays u p to 600 eggs 'at asitting', has a
lives onripefruits,whichplainlyaretemporary generation time of a little over three weeks, and large
habitats. Ifa fruit fly should attaln adulthood,I t simply populatlonoutbreaks occur ontheyounggrowth
migrates to another rlpe frult. Under these circum- of grasses, including m a w (Zed muyr). Some birds are
100 POPULATIONS

r-strategists.Thequail (Cotrtrnrx I-oturnix cot/trnrx) IS a habitatsand reproduce only by seeds (Hilligardt

case in point
(Puigcerver et ul. 1992). 1993).Trifolirrmpullese.rt~.sgrows on alpine scr'ces, often
in morainic areas; Trifolirrtn tbulii is predomlnantly
found in subalpineandalpinepastures. Trifoli/rn/
The K-strategists
tbalii, a plant of more competitor-Influenced habltats,
K-strategists occupy stable habitats. They maintain is a K-strateglst; Tr~ulirrrt/p~llesc.en.s
has characterIstlcs
populatlonsatstable levels, and areintensely of an Invasive, pioneer r-strategist.It has less devel-
competitive.They areoftenselected for large slze. oped vegetativestructures,hlgher seed productlon,
Many vertebrates are K-strateglsts.They arecharac- hlgher seed-bank reserves, smaller seeds, andhigher
teristicallylarge, long-lived, very competltive, have rates of seedlingmortality
low birthanddeath rates, and Invest muchtime
and effort in raising offspring. Their low growth rate
Competitors, stress-tolerators,and
helps to avoid thehabitatdegradationthat would
ruderals
follow an overshooting of the carryingcapacity. ~~

AnlmalandplantK-strategistsaliketendto invest Some populations are finely adjusted to habltats that

much energy in defence mechanlsms.Animal K-
strategists often have small litters or clutchesand
parents are very careful wlth their young. An extreme
example is the wanderlng albatross (Drornediarxuluns).
This bird breeds in alternate years (when successful),
lays just one egg, and is immature for nine to eleven
years, the longest period of irnmaturlty for any blrd
(see p. 94).
Not all vertebrates are K-strategists. The nomadic
Australian zebra finch (Tuenzopygiu emtunatis) is
r-selected, and the blue tit (Purrrs mrrdu.r), with the
largest recorded clutch size for a passerine blrd, is an
opportunlst. If their populatlons should fall drama-
tically, thenthebirth ratewilloften Increase to
bringthepopulatlonbounclng back to Its stable
level. This happens by increaslng the litter or clutch
size. Closely related species may displaydifferent
population strategies. Two trefoil species (Trifoliutn)
in Europe - the pale trefoil ( T r i f o h t i PulfescenJ)
and Thal's trefoil (Trifolirtm thalii) - prefer distinct
vary considerably In time.Wheretemporalhabitat
variationsare pronounced,populations may experl-
ence adverse conditions or stress. Athree-strategy
model,whichaddsastress-toleratlonstrategyto
competitiveandopportunisticstrategies, has been
applied to plants (Grlme 1977; Grime et ul. 1988).
Externalenvironmental factors that affect plants
fall into twochlef categories - stress and disturbance.
Stress lnvolves all factors that restrict photosynthesls
(rnalnly shortages of light, water, or nutrients; and
suboptimal
temperatures).
Disturbance involves
partial or total destruction of plant biomass arrslng
from herbivores, pathogens, and humans (trampling,
mowing, harvesting, and ploughing), andfrom wlnd-
damage, frosting, droughting, soil erosion, and fire.
Combinlng high and low stress with high and low
disturbance yieldsfourpossible contingencles to
which vegetatlon adapts (Table4.4). Nonce that there
is no practlcablestrategy in plants for living in a
stressful and highly disturbed envlronrnent. This is

Table 4.4 Environmental contingencies and ecological strategies in plants

Disturbance intensiiy Stress intensiiy

~

low High

Low Competitors Stress-tolerators

High Ruderals (No practicable strategy)

Source: After Grime (1 977)

 POPULATIONS 101

Table 4.5 A key for identifying ecological strategies of herbaceous plants

Levels of dichotomous
Strategy key

I 2 5 3 4

Annual + Potentially + Floweringprecocious Ruderal

fast-growing
+ Flowering delayed Competitive-ruderal
+ Small and slow-growing Stress-tolerant-ruderal
Biennial 3 Potentially largeand fast-growing Competitive-ruderal
+ Small and slow-growing Stress-tolerant-ruderal
Perennial + Vernalgeophyte Stress-tolerant-ruderal
+ Not vernal + Rapid + Rapidproliferationand Competitive-ruderal
geophyte
leaf
turnover
fragmentation
of shoots
+ Shoots not + Shoots tall and Competitor
fragmenting
laterally extensive
rapidly
+ Shoots short or Competitive-
creeping stress-tolerant-ruderal
(C-S-R)
+ Slowleaf + Shootstall, or laterally Stress-tolerani
extensive,
turnover competitor
both or
+ Shootsshort and notlaterally Stress-tolerator
extensive

Source: After a diagram in Grime et a/. (1 9 8 8 )

because contlnuousand severe stressin highlydis- may be displayedon a triangular diagram(Figure

turbed habitats prevents a sufficiently swift recovery 4.8). The plants assoclated wlth each strategy share
or re-establishment of the vegetation. (Some humans the sameadaptive characteristics. The following
- teachers come to mlnd - manage to survive under examples are all Brltish plants:
those conditlons.) The remaming three environmental
contingenciesappear t o have produced three basic 1 Competitorsexploit low stress and low distur-
ecologlcal'strategies In plants - competitors, stress- bance envlronments.The rosebay willow-herb
tolerators, and ruderals. Competltors correspond to (Chamerron anpstijiditm) occurs In a wlde varlety
K-strateglsts,and ruderals tor-strateglsts. Stress- of undisturbed habltats. It is notably common In
tolerators are an additlonal group whose strategy I S a fertile, derelict
environments (urban clearance
response tothetemporalvarlabilityand adversity sites, cinders, building rubble, m m n g and quarry
of the physrcal envlronment. In additlon, there are waste, andother spoilheaps). I t I S abundant in
four intermediate ecological strategies - competltlve woodland glades, wood marglns, scrub, and young
ruderal, stress-tolerant competltor, stress-tolerant plantations; it I S wldespread in open habltats such
ruderal, and competitor-stress-tolerator-ruderal strategy as walls, cliffs and rock outcrops, waysrdes and
(or C-S-R). Strategies of different herbaceous plants waste ground. and river banks; and seedlings are
may be found uslng a dichotomous key (Table 4.5). sometimes found in wetland (but not submerged
The primary and intermediate ecological strategies areas), arable fields, and pastures.
102 POPULATIONS

2 Stress-tolerators exploit
high stress, low distur- 4 Competitive ruderals
exploit low stress,
bance, low competltionenvlronments.Theheathmediumcompetltmn,andmediumdisturbance
grass (Dunthonu decu?nbens) favours mountam environments.
The policeman’s helmet (Imaputmv
grassland. It I S also found in pastures andheath- glundrtlifrru) is abundant on river banks,shaded
land,and occasionally In road verges, grassy paths,marshland,anddisturbed,lightly shaded areas in
scree slopes, rock outcrops,
and
wetland. woodland and scrub.
3 Ruderalsexploithighdisturbance, low stress, and 5 Stress-tolerant competitors exploitmedium
low competitionenvironments.Theshepherd’scompetmon,medium stress, and low disturbance
purse ( C U ~ J ~hnrsu-pustori.r)
/U lives on disturbed,envlronments.The malefern (Dryuptens jlix-rnus)
fertile ground. It is especially abundant as a weed is restricted totwo very different habitats -wood-
on arableland and in gardens,and occurs In a land habitatsand skeletal habitats(mainly cliffs
range of disturbed artificial habitats such as demo- and walls, but also quarry spoil, rock outcrops and
litionsites,paths, spoil heaps, andmanure heaps. screes, rwer banks, andsomecmdertips).

1 Competitor
Rosebay willow-herb (Chamenon angustifolium)
Stinglng nettle (Urtica dioica)
Reed grass (Phalans arundinacea)

4 Competitive ruderal 5 Stress-tolerantcompetitor

Policeman’s helmet (Impatiens glandulifera) Male fern (Dryoptens filix-mas)
Creeplng buttercup (Ranunculus repens) Dog’s mercury (Mercurralis perennfs)
Coltsfoot (Jussilago farfara) Bilberry (Vaccfnwm myrtillus)

3 Ruderal 2 Stress-tolerator
Shepherd’s purse
(Capsella bursa-pastons)
Groundsel
(Seneoo vulgarrs)
Small nettle
(Urtica urens) 0 (Sanfcula europaea)
0
100 $0
Disturbance (per cent)

6 Stress-tolerantruderal 7 ’C-S-R’ strategist

Carline thistle (Carlina vulgaris) Yorkshire fog (Holcus lanatus)
Hard poa (Desmazerra rrgfda) Cat‘s ear (Hypochaerfs radicata)
Purging flax ( h u m catharticum) Common sorrel (Rumex acetosa)

F i g w e 4.8 Ecological strategles In plants. The triangular diagram I S used to define v m o u s mlxes of three baslc strategles
- competitor, stress tolerator, and ruderal. Pure cornpetltors lie at the top corner of the triangle, pure stress-tolerators at
the bottom rlght corner, and pure ruderals at the bottom left corner. Intermediate strategles are shown. Example spec~es
are all Brirish.
Sor/r.rr: After Grime rt d . (19x8)

Stress-tolerantruderalsexploitmediumdis-
turbance,medium stress, and low competltlon
environments. The carline thistle (Carlitla t d g u r ~ s )
occursin areas of discontinuousturf associated
with calcareous pastures, rock outcrops, lead-mine
heaps, and derelict wasteland. It I S also frequent
on sand dunes.
Competitor-stress-tolerator-ruderal (C-S-R)
strategists occupy a middle-of-the-road posltion
(mediumcompetition,medium stress, medium
disturbance) that enables them to live In a wide
range of conditions.TheYorkshire fog (Hob
lanatw) IS recorded as seedlings in every type of
habitat. I t is most abundant inmeadows and
pastures,and is prominent In spoilheaps,waste
ground, grassverges, paths,and inhedgerows.
I t is less common on stream banks, arable land,
marsh ground, and In skeletal habltats such
as walls and rock outcrops. It occurs In low fre-
quency in scruband woodland clearings,and I S
found also In grassy habitats near the sea and on
rnountalns.
Differentlife-forms tendto have a narrowrange
of ecological strategies. Trees and shrubs centre on
polnt
a between the competitor-stress-tolerator-
ruderal and competitive-ruderal positlons. Herbs have
a wide range of strategies, though competitor-stress-
tolerator-ruderal is common. Bryophytes (mosses and
liverworts)aremainlyruderals and stress-tolerator-
ruderals. Llchens are malnly stress-tolerator-ruderals
and stress-tolerators.
Migration strategies
Migration rates vary conslderablybetweenspecles.
In plants, three baslc mlgratlon strateb'rles seem to
exist and are analogous to ruderal, opportunist, and
competitive population strategles.
Fugitive species
These are the 'weeds' of the plant kingdom. They
colonize temporary,disturbedhabltats, reproduce
rapldly, and soon depart before the habitat disappears
POPULATIONS 103
or before competitionwithotherorganisms over-
whelmsthem.Thecommondandelion (Taraxmutl
rfficinale) is a fine example. A fugitive-migration
strategist occupies a patchwork of habitat islands.
The tamarack (Larix laricina) is a fugitive-strategist
(Delcourt and Delcourt 1987: 319-22). Its changlng
distribution mirrors
the
availability of wetland
habitats, and any snapshot of its distribution shows
low dominance in most areas, with dominance hot-
spots in local wetland sites (e.g. Payette 1993).
Opportunist species
Fast migrators spread rapidly, pushing forwards
along a steep migration front, butfailing to mamtaln
large populations in Its wake.Spruce (PilzL7) I S an
r-migration strategist and displays these character-
istic(DelcourtandDelcourt 1987: 306-12). With
the retreat of the Laurentide Ice Sheet, spruce spread
rapldly onto the newly exposed ground. The rate of
migration reached 165 m/yraround 12,000-1O,Oo0
years ago. The spruce's northward advance was
halted only by the Ice. Behinditsmigratlonfront,
populationsdiminished as climateswarmed. Black
spruce (Piaa marzana) was the first conifer species to
invade northernQuebecimmediately after the ice
had melted 6,000 years ago(DespontsandPayette
1993).
Equilibrium species
Slow mlgrators rlse more slowly to domlnance after
an lnltlal invasion, the highest values of dominance
lying well behind the 'front lines' and reflectingan
unhurried build-up of populatlons. Oak (Qrwws) I S a
K-migration strateglst. It was a minor constituentof
late glaclal forests in eastern North America. It did
reach the Great Lakes in late glaclal times,but Its
r s e to dominance was a slow process that reached a
ceiling in the Holoceneepoch In the region now
occupled by the eastern deciduous forest (Delcourt
and Delcourt 1987: 313-16).
104 POPULATIONS
P L A Y I N G WITH N U M B E R S :
P O P U L A T I O NE X P L O I T A T I O NA N D
CONTROL
Exploiting populations
The commercial exploitation of marine mammals,
fish, big game, and blrds has caused the extinction
of several specles and the near-extinction of several
others. The cases of the passenger pigeon, great auk,
northern fur seal, and the Amerlcan bison illustrate
t h ~ polnt.
s and puffins. It used to nest aroundtheNorth
Passenger pigeon
These graceful blrds once flourished in North
America. O n 1September1914,the last passenger
pigeon (ktopz.rtt.r tt/qyaturz/u),called ‘Martha’, died in
the Cinclnnati Zoolog~cal Garden. Therapid extinc-
tion of such a flourishing population resulted from
three misfortunate circumstances. First, the pigeons
were very tastywhen roasted or stewed. Second,
they migrated In hugeanddense flocks, perhaps
containmg 2 billion birds, that darkened the sky as
they passed overhead. Third, they gathered in vast
numbers,perhaps several hundredmillionbirds,to
roost and to nest. A colony In Michigan was estimated
t o be 45 km long and an average of around 5.5 km
wide. Professional hunters - pigeoners - formed large
groups to trapandkillthepigeons.Nesting trees
were felled to collect thesquabs(young, unfledged
pigeons). The clearrng of forest for farmland and
theexpans~on of the railroad madehugetracts of
the plgeon’srange accessible andthe persecution
intensified. By about 1850, several thousand people
were employed in catchlng and marketing passenger
plgeons. Every means imag~nablewas used to obtaln
the tasty bird. Specla1 firearms,cannons, and fore-
runners to themachrne gun were built. In 1855 alone,
one New York handler had a daily turnover of 18,000
plgeons. In 1869, 7.5 million birds were captured at
one spot. In 1879,1 billion birds were captured in the
state of Michigan.Theharvesting rate disrupted
breeding and the population S I X began to plummet.
Even by 1870, large breeding assemblies were
confined to the Great Lake States, and the northern
end of the pigeon’s former range. The last nest was
found In 1894. The last bird was seen In the wild in
1899.
Auks
The ‘penguin of the Arctic’, the great auk ( P i n p i n ~ l s
ztnpennzs), was an early casualty of uncontrolled har-
vesting.This flightless bird was the largest of the
Alc~dae,whichfamilyIncludes guillemots,murres,
AtlanticOcean, in north-easternNorth Amerlca,
Greenland, Iceland, andthenorth-westernBritlsh
Isles (Figure4.7).The last palr died in Iceland in
1844. A few museum speclmens survive.
The razorbill ( A h tor&) is the largest living auk.
For many years, this species was killed for its feathers,
and by fishermen for use as bait. This exploltatwn
caused the species todecline.The razorbill is now
protected and I S stagmg a comeback, so it may avoid
the fate of its close relative, the great auk.
Fur seals
Millions of northern fur seals (Ca//orhznm ursinrrs)
used to breed on the Pribilof Islands, in the Bering
Sea, Alaska.Between 1908 and 1910, Japanese seal
hunters slaughtered 4 million of them. The Fur Seal
Treaty of 191 1was signed by Japan, Russia, Canada,
and the United States. Thlswas the first International
agreement to protect marineresources. The signatories
agreed rates of harvesting for pelts. The happy result
was thatthenorthern fur seal populatlon bounced
back. New colonies occaslonally appear. Northern fur
seal pups were first observed on Bogoslof Island, In
thesouth-eastBering Sea, in 1980(Loughlinand
Miller 1989). By 1788 the populatlon had grown to
more than 400 individuals, including 80 pups, 159
adult females, 22 terr~torialmales, and 188 ~uvenile
males. Some anlmals origlnated from rookeries of the
CommanderIsland;others were probablyfrom the
Pribilof Islands.
 POPULATIONS 105

Figure 4.9 The great auk‘s (Pingumus rmpmnzs) former distributlon. Black dots areformer nestmg grounds. The whlte
clrcle marks the sxe where the last two birds died In 1844.
Soutre: Map after Ziswiler (1967); ptcture from Saunders (1889)

American bison encouraged buffalo shoot-outs from the trains. Thus

was perpetrated a senseless and bloodthirsty slaugh-
Humans have hunted terrestrial, as well as marme, ter. Tens of thousands of animals were shot and left
mammal species to extinctron or near extlnctlon. A to rot alongslde the tracks. By the 1890s, only a few
spectacular,
shameful,
if example
concernsthe hundred buffaloes werestillaliveintheUnlted
American bisonor ‘buffalo’(Bison bison) (Figure 4.10). States, and a few hundred more of the wood buffalo
In 1700, the buffalo populatron on the Great Plains varlety In Canada. In 1902, the population was less
was around 60 million. Native Indians had hunted than a hundred. Happily, buffalo breed well under
this population sustainably for thousands of years. A anlmal husbandry. Thanks to William T. Hornaday
newera of buffalo huntingstartedinthe1860s, of the New York Zoologlcal Society, the American
when the Union Paclfic Railway was opened. At first, BisonSociety(foundedin1905),and theUnited
speclal hunting partieswerehired tosupplythe StatesandCanadiangovernments,the specles was
railway workerswithmeat.William ‘BuffaloBill’ saved. There was a vigorous programme of captive
Cody rose to fame at this time for having killed 250 breedingandreleasrng of anlmalsontoprotected
day! The tracks of the UnionPacific ranges, such as that In Yellowstone Park. The bison
buffalo in a single
cut rightacross the heart of the buffalo country. Once has nowrecovered to a population size of tens of
the railway was operatmg,the railway company thousands and has been Introduced into Alaska.
106 POPULATIONS

Controlling populations Swedish beavers

Muchconservationeffort IS currentlyexpendedon
reintroducing species to least
at someof their former
domam, and keepmg successfully introduced specles
under control. These two aspects of populatlon man-
agement are exemplifiedby beaver reintroductlons in
Sweden and by managmg feral goatsIn New Zealand
and feral cats on Marlon Island.
The European beaver (Custwjber) is Europe's largest
rodent.It livesbesideshores of watercoursesand
lakes,inexcavatedburrows or constructedlodges.
The beaver was heavily hunted In Sweden during the
nmeteenth century (Bjarvall and Ullstrom 1986:77).
By the time it received protection In Sweden, it was
too late - the last animals had been shot two years
earlier.In 1922,reintroductions were begunusing
animals captured In southern Norway. The popula-
tlon grew to about400 animals in 1939 and 2,000 in
1961.During the 1960s the population trlpled and

Ir
bison American v"r
, 1, Bison
bison I ~

yl

w 1875
r-"J 1850
..........
..................
..........
..........

Figure 4.10 The shrinking range of the American bison (Bison bzson).
Source: After Ziswiler (1967)

increased to 40,000 animals by 1980. Given such a
large increase, somehuntmg was permlttedfrom
1973.Thehunting appears to have been justified.
Therelntroduced beaver population in Sweden has
behaved as introducedungulatepopulations - It
‘exploded’ andthenstartedtodecline(Hartman
1994).In two studyareas, the populatlon growth-rate
turned negative after 34 and 25 years and at densltles
of 0.25and0.20 colonles per km’, respectively.
Management policy for an Irruptivespec~esshould
probably allow hunting during the Irruptive phase.
Huntlng keeps the irruptive population In check and
so maintalns food resources and avoids uncontrolled
population decline.
N e w Zealand goats
Feral goats (Capvu hrvc-lr.r) live In about I I per cent of
New Zealand, mostly on land reserved for conserva-
tion of the indigenous blota. They were introduced
from Europe before 1850. They were recognized as a
threatto native vegetation from the1890sonward
and soon attained the rank of a pest. The effect of
their browsing is most noticeable on islands, where
subtropical forest has insome cases been converted
to rankgrassland (Rudge 1990). Uncontrolled goat
densities areusually less than 1 per ha,but have
reached 10 perhain one area (Parkes1993).The
total population is at least 300,000. Goats are highly
selective in what they eat, but switch their diets as
the more palatable food species are eliminated from
their habitat.
Unharvested feral populations In NewZealand
form stably dynamic K-strategist relationships with
their resources. Their impacts are thus chronic, but
therelationship can be manipulatedto favour the
resource if management can be applied ‘chronically’,
that is, as sustainedcontrol.TheDepartment of
Conservation has attemptedto organize itscontrol
actions in the highest priority areas either to eradi-
cate goats where possible or to sustain control where
eradicatlon is not presentlypossible. In 1992-3,
6,160 hunter-days of ground-control effort, and 230
hours of helicopterhunting effort, were expended
against feral goats, and 10.2 km of goat fencing was
POPULATIONS 107
constructed(thoughgoats arenotoriouslyhard to
fence In). If the present effort is maintained through
to the first decade of the new millennium, about half
the feral goatpopulatlons now on the conservation
estate areexpected to be eradicated or controlled.
However, goats are a valuable farming resource, and
this may assure the survival of feral populations in
hobby farms and hill country farms bordering forests
(Rudge 1990).
Marion Island cats
MarionIsland is the largest of the PrlnceEdward
Islands and lies In the southern Indian Ocean, about
midwaybetween Antarcticaand Madagascar. It is
a volcanic island wlthan area is 29,000 ha that
supports a tundrabiota.Domestic cats (Felis catu1)
were introduced to Marion Island as pets in 1949. By
1975, the population was about 2,139 and had r m n
to 3,405 in 1977. As on other sub-Antarctic islands
wherecats were introduced,the feral cats played
havoc with the local bird populatlons. They probably
caused the local extinction by 1965 of the common
divlngpetrel (Pr1ecanotde.r unnutrzx), and reduced
thepopulatlonsandbreeding successes of wmter-
breeding great-winged petrels (Pterodroma ~nacroptera)
and grey petrels(Procellaria cinerea). Control of the cat
population was begun in 1977.The viral disease
feline panleucopaenia was chosen as the most efficient
and cost-effective method. The population fell from
3,405cats in 1977 to 615 cats in1982.However,
thedecline slowed downand evidence suggested
that biological control was becoming ineffective. A
full-scale hunting effort was launched in the southern
spring of 1986 (Bloomer and Bester 1992). The aim
was to eradicate the cat populationfrom the island. In
14,725 hours of hunting, 872cats were shot dead and
80 were trapped.Thenumber of cats sighted per
hour of night hunting decreased, which was strong
indication that the population had declined. But, by
the end of the third season, it became evident that
hunting alone was no longer removing enough ani-
mals to sustain the population decline, andtrapping
was incorporatedintotheeradicationprogramme.
Trappingwithbaitedwalk-in cage traps has a
108 POPULATIONS

drawback: other specles,such as sub-Antarctic skuas E S S A Y QUE ST I O N S

(Catharactuanturtncx), also become ensnared. How-
ever, a small loss of non-target species has to be offset 1 Why do some populations irrupt
agamstthelong-term benefits of the cat trappng and crash?
programme to the bird populatlon.
2 What are the advantages and
disadvantages of the different
SUMMARY strategies? survival population

Populations are Interbreeding groups of Individuals 3 Why do some populations need

(demes).Unconstramed population growth describes controlling?
a rising exponential curve; when constramed, I t
describes a logistlc curve, or may behave chaotically.
Populatlon irrqmons and crashes are common. Age F U R T H E RR E A D I N G
and sex are Importantcomponents of population
growth. Theyare summarized In life tablesand Hanskl, I. and Gilpin, M. E. (1997) Metapoprdatron
survlvorship curves, and they are studiedusing Brology: Erology. Genetra. and Evulrrrron, NewYork:
cohort-survival models. Metapopulations are defined Academlc Press.
according to the geographlcal locatlon of indivlduals
withln a population - whether they form one large Kormondy, E. J. (1996) Concepts ufE'c.ology, 4th edn,
interactinggroup, or whether they live in several Englewood Cliffs, NJ: Prentice Hall.
relatively Isolated subpopulations, for example.
Recent work on the conservation of threatened popu- Kruuk, H. (1995) Wild Otters: Predution and
lationsdraws heavily on metapopulationtheory. Pop~lattuns, Oxford:Oxford University Press.
Populatlonsadopt varlous strateglestomaxlmize
their chances of survival. Opportunists (r-strategists) Perry, J. H., W o ~ w o d ,I. P.,Smith, R . H.,and
and competltors (K-strategrsts) are two baslc groups. Morse, D. (1 997) Chaos rn Real Datu: Ana1ysr.r on Non-
Stress-toleratorsform another strategic option. The ~ ~ Short EcologrLxl Time
linear D y n a v ~ rfrom Series,
three basic migration strategles are fugitives, oppor- London: Chapman & Hall.
tunlsts,andequilibrists.Wildpopulations suffer
heavily at the hands of humans. Some species were Taylor, V. J.andDunstone,N. (eds) (1996) The
huntedtoextinction.Others were huntedto near Exploltatrorl of hlatnrnal Po~tduttons,London: Chapman
extinction butmanaged to recover. Population control & Hall.
is often necessary to prevent damage to agrlcultural
resources andenvironmentaldegradation.Control Tuljapurkar, S. and Caswell, H.(1997) Strr/r.trwed-
involves culling or eradicating introductlons, such as Poprdatron Modeh, New York: Chapman & Hall.
goats andcats. It may also may involve re-establishing
a specles over vacated parts of Its former range.
 5

INTERACTINGPOPULATIONS
Populations interact with one another. This chapter covers:

co-operating
populations
competing
populations
plant-eating
populations
flesh-eating
populations
biological
population
control

Twopopulations of thesame species, living In theInteraction(thelnteractlon I S notobligatory). Some

same area, may or may not affect each other. Inter- small blrds rldeon the backs of waterbuffalo - the
actlons betweenpairs of populationstake several birdsobtain food and In dolng so rld the buffalo of
forms, depending on whetherthe influences are Insect pests. Some birds plck between theteeth
beneficlal o r detrlmental t o the parties concerned of crocodiles - the blrds get a meal and the crocodile
(Table 5.1). Inmutually beneficial relatlonships,gets a free dentalhygiene sesslon.
both populations gain; in mutually detrlmental rela-
tionships, both lose. Other permutatlons arepossible.
Animal-animal protocooperation
One .population
. may galn and the other lose, one may
galnandtheother be unaffected; one may lose andAremarkableexample of protocooperation I S the
other
the be unaffected. partnership between
an Afrlcan
greater
the
bird,
honey-guide (Inclimtur rnrjicator), and a mammal, the
honey badger or ratel (Mellivuru c-upensis) (Plate 5.1).
LIVING TOGETHER: C O - O P E R A T I O N
Thehoney-gulde seeks out a beehlve, thenflitters
Four types of lnteractlons between populatlons or around making a chattering cry to attract the attention
specles Favour peaceful coexistence - neutralism, of a honey badger, or any other interested mammal,
protocooperatlon, mutualism,and
commensalism. Including humans. The honey badgerrips open the
Neutralism Involves no lnteractlon berween two hlve and feeds upon honey and bee larvae. After
populatlons. I t I S probably very rare, because there the honey badger has had its fill, the bird feeds. Its
arelikely to be indirectinteractions between all digestive system can even cope wlth the nest-chamber
populatlons In a glven ecosystem. The other forms of wax, which is broken down by symblotic wax-
interaction are common. digesting bacteria living in its intestines. In captivlty,
Indivltluals have been kept alive for up to 32 days
feeding exclusively on wax! The honey-guide is adept
Protocooperation
at finding beehlves but is incapable of openlng them;
Protocooperation involves both populations benefit- the honey badger IS good at opening beehives but is
ing one another in some way from thelr Interaction. notgoodatfindingthem.The protocooperation
However, neither population’s survival depends on the clearly profits both specles.
1 10 INTERACTING
POPULATIONS

Table 5. I Interactionsbetweenpopulationpairs

Kind of interaction Comments

Species

A B

Neutralism N o effect N o effect The populations do not affect one another

Protocooperation Gains Gains Both populations gain from their interaction,
but the interaction is not obligatory
Mutualism Gains Gains Both populations gain from their interaction,
and the interaction is obligatory (they
cannot survive without it)
Commensalism Gains No effect
Population Acommensal)
(the gains;
population B (the host) is unaffected
Competition TheLoses
Loses populations inhibit
another
one
Amensalism Loses No effect Population A is inhibited; population B is
unaffected
Predation Gains Loses Population A (the predator) kills and eats
members of population B (the prey)

Plant-plant protocooperation carry their pollen tootherplantsandtheir seeds

Protocooperation seems to occur among plants. Two
species of bog moss, Sphagnum magellan~c~rnrand
Sphagnumpapillosum, were grownunder laboratory
conditions (Li et a / . 1992). Underdryconditions,
Sphagnum magellanimm outcompeted S p h a g n ~ npapil-
~ into new plants from the droppmgs. Mammals and
los~onfor water, largely because it is better designedfor
transporting and storing water. Sphagnmzmagellan-
iczcm is a drought-avolder and prefers peat hummocks,
while Sphagnm papillostrm is a drought-tolerator and
prefers lower sites on a hummock. However,Spbagntrm
nlagellanirrrm benefits Sphagnrrm papillosrrm growing at
higherhummockpositions by encouraging lateral
flow of water. Both species grow better when mixed,
suggesting a degree of protocooperation.
Animal-plant protocooperation
Plants often form co-operatwepartnershipswith
anlmals. The most common examples are assoc1atlons
between plants and pollinators, and between plants
and seed dispersers. Plants arerooted tothe place
that they grow so they must employ a go-between to
toother sltes. Colourful flowers with nectar and
brightly coloured fruit have presumably evolved to
attractanimals as potentlal delivery agents. Some
herblvores eatfruits,the seeds inwhich may pass
through the herbivore intestines unharmed and grow
blrds carry seeds in thelr fur and feathers. The sheep
carries agarden-centre-full of plant seeds around
in its fleece (p. 60). Birds, bees, andbutterflies are
the
commonest pollinators. Some mammals are
pollinators,includinghumans, nectar-feeding bats,
and the slender-nosed honey possum or noolbender
(Tarsipes rostratrrs). The noolbender is adiminutive
Australian marsupial thatfeeds on nectar, pollen and,
tosomeextent,small Insects that live in flowers
(Colour plate 7).
An unusual example of plant-animal co-operation
is the alliancebetween some species of Acacia and
epiphytic ants. Species of acacla that normally sup-
port ant colonies are highly palatable to herbivorous
insects; those that do not normally house an ant fauna
to ward off herblvores are less palatable. Acacias
guarded by ant gangs produce nectaries and swollen
 INTERACTING POPULATIONS 111

Plate 5.1 Ratel or honey badger (Mellivom capnsis)

Photograph by Pat Morris

thorns that attract and benefit the ants. The plants butterflies (Heliconius) are confined to the Americas.
put matter and energy into attractlng ants that will Therearesomefortyspecles,most of whichare
protect their leaves, rather than into chemical war- restricted to rain forest. In different parts of South
fare. This antiherbivore ployIS broad-based, since the America, the various species tend to look much alike
ants fiercely attack a wrde rangeof herbivores. andformMullerianmimrcryrings. I t isnottoo
difficult to see theadvantage of suchbiological
photocopymg. Tenderfoot
predatorspresumably
Mimics learntoavoidunsavoury prey by trialanderror,
Miillerian mimicry (namedafterFrrtzMuller, a killing and eating at random. If distasteful species
nineteenth-centuryGermanzoologist)occurswhen vaned widely in appearance, then the predators would
two or more populations of distasteful ot dangerous be forced to kill many of each before learning those
species come to look slmilar. An example IS bees and to avord. However,whentheunsavourypreyare
wasps, which are normally banded with yellow and coloured in the same way, then the predator qulckly
black strlpes. In Trinidad, the unpalatable butterflies learns to avoid one basic pattern. Its sample snacks
Hirsutzs mgara (from the familyIthomiidae)and are therefore spread out over many prey speaes. By
L y c m ceres (from the family Danaldae) resemble one resembling one another, unsavoury prey populations
another(Figure 5.1). The polsonouspasslon-flower keep their losses by predatlon to a minimum.
1 12 INTERACTING
POPULATIONS
Hirsutis megara
Fiptre 5 . I The unpalatable Trlnidadian butterflies N/r.rtti/r r r w , q ~ w ' t (from the family Ithomiidae) and Lprra L-rres (from
the family Danaidae) resemble one another - they are Mullerian mlmics.
Sortrce: From Brower ( 1969)
Batesianmimicry,named afterthenineteenth-
century
Englishnaturalist
HenryWalter Bates,
occurs when a palatable animal comes to look like an
unpalatableone.Theharmlessdronefly (Erzstulis
tenax) has warning coloration much like a honey bee's
(Apn ttzellijiia). The advantageinsuchmimicry is
avoldance by would-be predators mistaking it for a
bee wlth a sting In Its tail. The dronefly, and other
mimics, is thus a 'sheep In wolfs clothing', bluffing
Its way through life on the strength of masquerade.
Batesian mimlcry is q u t e common In theanlmal
kingdom.Many speclesofharmlesssnakes mimlc
polsonoussnakes - harmlessandpoisonous coral
snakesinCentral Americaare so similarthatonly
anexpertcantellthemapart.Batesianmlmicry IS
disadvantageous tothe poisonous'model' because
predators will eat harmless mimics andso take longer
to learn toavoldthemodel'swarningcolours. So,
technlcally speaking, I t I S a brand of predatlon.
Mutualism
In a mutualistic relationshlp, both populations ben-
efit frominteracting.However,theyaredependent
on their interaction to survive. Mutualism I S far less
common than protocooperation, of which I t is a more
extremeform.It is the evolutionaryequivalent of
'puttlng all your eggs in one basket' - the reliance
ceres Lycorea
of
one specles onanother is complete. Certain
Australiantermltes, for example,cannotproduce
enzymes to digest thecellulose in wood. They exploit
wood as B food source by harbouring a populacion of
protozoans (Mpvtrithir pcrruhxu) capable of making
cellulose-digesting enzymes in their guts. The proto-
zoans are intestlnal endosymbionts. Nelther the
termltes n o r the protozoanswouldsurvlvewlthout
the other. One generaclon of termites passes on the
protozoans to the next by exchanplngintestinal
contents.
Lichens are thought to depend on amutualistic
alliance becween a fungus and an alga - the fungus
provides the shell, the alga the photosynthetic power
house.However,thealgae In somelichenscanbe
grown wlthout thelr fungal host, so the relationshlp
may In some cases be protocooperation.
Mutualism I S also known in marine environments.
Two encrustlng sponges (S/ll,rrrtes rul,r/r.s and S/rberitrs
//rrid/ls)protect the queen scallop (Chlrtty opert-uluvis)
frompredation by a starfish, the common crossfish
(Astevru.~rubens) (Pond 1992). Theyprobably do so
by maklng it difficult for the common crossfish to
g r l p thescallopwiththelrtube-like feet. and by
excluding other organisms settling on the scallop,so
hlndering its mobility. The sponge benefits from the
associatlon by belng afforded protectlonfromthe
shell-less nudibranch-gastropodpredator ArLhrdoris
 INTERACTING POPULATIONS 113

pseudoav~/rs,andmoregenerally by beingcarriedto chewing upon solid materials, while the mosquitoes

favourable locations. filter-feed ontlnybltsthat break off thedecaying
matter and on bacterla. Mosquitoes eating particles
has no effect on the midges, but midges feeding on
Commensalism solid material creates a supply of organlc fragments
Commensalism occurswhenonepopulation (the and, by increasingthe surfaceareaof decaying
commensal) benefitsfrom an Interactionand the matter, bacteria.
other population (the host) is unharmed. Bromeliads
and orchlds grow as epiphytes on trees wlthout doing
Mynas and kingcrows
any harm. The house mouse ( A l ~ tt/t/.rcu///.r
s c/ome.rtiu/s)
has lived commensallywlthhumansslncethe first In
India,
twoblrd species, the
common
myna
permanent settlements were established In the Fertile (Arvzdothrr/(srvirtrs) and the jungle myna (Acridotherru
Crescent(Boursot et a / . 1993). The housemouse f j l s c ~ s )forage
, in pure and mlxed flocks on fallow land
benefits fromthe food andshelterinadvertently (Veena and Lokesha 1993). Another bird species, the
provided by humans,whilethehumans suffer no klng-crow (Drcr/wus t/mmzrctl.r), a drongo, feeds on
adverse effects. Four other examples are as follows. Insects disturbed by the foraglng mynas. King-crows
tend t o consort wlth larger myna flocks, comprising
Cattle egrets and cattle twenty or moreIndivlduals. Thekmg-crowsand
mynaseatdifferentfoods, s o they do not compete.
Cattleegrets ( B N ~ N I Mihrs),
S whicharenatlve to The klng-crows benefit from the assoclation, whereas
Afrlca and Asla, follow cattlegranng In theSun, the mynas are unaffected - a case of commensalism.
pouncingoncrickets,grasshoppers, flies, beetles,
lizards, and frogs flushed from the grass as the cattle
approach (Colour plate 8). The cattle are unaffected River otters and beavers
by the blrds, while the birds appear to benefit - they
In the boreal forest of north-east Alberta, Canada, a
feed faster andmore efficiently when associated
partlal
commensalism has evolved between rlver
wlththecattle.Interestingly,thecattleegret has
otters (Lutra ~ut~acfensts)
and beavers (Castor 1unarlensr.r)
expanded its range, successfully colonizing Guyana,
(Reid et a / . 1988; Colour plate 9). In winter, otters
South America, in the 1930s. It filled a niche created
dig passages through the beaver dams to guarantee
by extensiveforestclearance. Without largeherbi-
vores to stir u p its meal, it has taken to feeding in the underwater access to adjacent water bodies, which are
company of domestlc livestock, which also disturbs frozen over. The passages lead to a reduction in pond
insects and other prey living in grass water-level, whlch may Increase the otters’ access to
air under the ice, and t o concentrate the fishes upon
whichtheotters feed. Beavers repairthe passage
Midge and mosquito larvae in pitcher-plant during timesof open water, so otters benefit from the
pools creation andmaintenance offish habltats by the
InNewfoundland,Canada,thecarnivorouspltcher beavers, who themselves are unaffected by the otters’
plant, Sarracenia pwpurea, has water-filled leaves activltles.
holding decaying invertebrate carcasses (S. B. Heard
1994). Larvae of a midge (Metriocxmm knabi) and a
mosqulto (Wycottzyia mithii) feed on the carcasses. S T A Y I N G APART: C O M P E T I T I O N
The two Insect populations are lirnlted by the carcass
supply, but the use the carcasses at different stages Whilesome speclesco-operatewithoneanother,
of decay and live commensally. The midges feed by others compete for food o r territory.
114 I N T E R A C T I N GP O P U L A T I O N S

Competing for resources Scramble competition

Competition is a form of popt~lat~on interaction In
Competitlon between populations occursin two ways
whlch both populations suffer. It occurs when two o r
-- scramble competition and contest competition.
morepopulatlonscompete for a common resource,
First, scramble, exploitation, or resource competltion
chiefly food or terrltory, thatI S limited. It may engage
occurs when two or more populatlons use the same
members of the same species (intraspecific compe-
resources (food or terrltory), and when those resources
tition) or members of two or more different species
are In short supply. This competition for resources
(interspecific competition). The limitingresource
I S Indirect. One population affects the other by
prevents one or both the species (or populatlon) from
recluclng the amount of food o r territory available to
growing.
the others.
classlc
A example of scramble competition
Competitive exclusion concerns the natlve red squirrel (.Yu/wtll w//prl.r) and
the Introducedgrey squ~rrel(.Yc/ur/lJ r~uro/incmsi.r)~n
The outcome of competlng for a limiting resource
England,Wales,andScotland(Figure5.2;Colour
I S not obvlous and depends upon a varlety of circum-
plate 1 0 ) . The red squ~rrelfavours coniferous forest,
stances. Itwould be temptlngtothlnkeitherthat
but also lives in dec~duouswoodland,particularly
both species may coex~st, thoughat reduced density,
beech woods. The grey squlrrel, a natlve of eastern
or else thatonespec~es maydisplacetheother.
North Amerlca, is adapted to life In broad-leaved
However, coexlstence I S exceedinglyunlikely. As a
forests, but I S adaptable and will colonlze coniferous
very strictrule,notwo spec~escan coexist onthe
forests. In Brltaln,
the red squirrel’s
rangehas
same limltlng resource. If they should try to do so,
progressively shrunkduringthetwentlethcentury
one specles will outcompete the other and cause its
asthegreysqu~rrel,introducedfromAmerlca In
extlncrion. Thls finding I S encapsulatedin Gause’s
the late nineteenth and early twentiethcenturies,
principle, named after Georgii Frantsevlch G ~ L I sIt~ .
hasextendeditsrange. A dramaticdeclinein red
I S also called the competitive exclusion principle
sqwrrelrangeoccurredfrom1920to1925, largely
(Hardin 1960). It was first establishedinmathe-
dueto disease following greatabundance(Shorten
maticalmodels by Alfred James Locka (1925)and
1954). The grey squirrel rapidly advanced Into the
Vito Volterra (1928), and later in laboratory experi-
areas relinqulshed by the red squirrel, and into areas
ments by Gause (1034).
around thelr introduction sltes. Up to around 1960,
Gause used culture
a medium to grow two
the red squlrrelstill had a few strongholdsinthe
Paratt/m/m specles - Purun/ecmm awe/iu and
southern parts of England,especially East Anglia and
Pururtzt.l-itm/cmdutmtt. Each population showed a
the New Forest. The grey squirrel has advanced into
logistic growth curvewhen cultured
separately.
these areas and replaced the red squirrel in much of
Whengrowntogether,theydisplayed a loglstic
East Anglia (H. R. Arnold 1993). Red squirrels can
growth curve for the first S I X t o e g h t days, but the
stillbefoundonthe Isleof Wight and Brownsea
Pururt/ecintt/c.rrtduttln/ populatlon slowly fell and the
Island In Pook Harbour, Dorset.
Purunteczunt u/m/iu population slowly rose, though
The success of the grey squirrel may lie primarily
it never attainedquite so hlgha levelas when it
In Its superlor adaptability to the herbwore niche at
was cultured separately.
Manyother
laboratory
canopy level in d e c l d ~ o u swoodland (C. B. Cox and
experimentsusingvariouscompetlngpopulatlons
Moore 1093: 61).However,onestudyhighlights
givecomparableresults. Competitors used include
the complexity of squirrel competltlon (Kenward and
protozoa, yeasts, hydras, Dupbttiu, gram beetles, frulc
Holm1993). Conservatlve
demographic
tralts
flies, and duckweed (Lrmm).
comblnedwlthfeedingcompetitioncouldexplain
the red squirrel’sreplacement by greysquirrels. In
 INTERACTING POPULATIONS 115

Red squirrel
Sciurus vulgaris

x *. ."... ... .......

....... 0
Grey squirrel
Sciurus carolinensrs

0
1 16 INTERACTING P O P U L A T I O N S

oak-hazel woods,grey squirrelforaging,density, on rlvals. Thenoddingthistle (Carduus nutans) I S

and productivity are related to oak (Quwzus spp.) and allelopathic, releaslng solubleinhibitorsthatdis-
acorn abundance.Incontrast, red squirrels forage
courage the growth of pasture grasses and legumes,
where hazels (Covybs avellana) are abundant;their at two phases of its development - at the early bolt-
relatively low density and
breeding ing phase when larger rosette leaves are decomposing
success are
related to hazelnut abundance. Red squirrels fail to and releasing solubleinhibitors,andatthe phase
exploit good acorn crops, although acorns are more when bolting plants are dying (Wardle et al. 1993).
abundant than hazels. However, in Scots pme (Prnus Moreover, noddingthistleseedlings seem to be
sylvestrrs) forests, their density and breeding stimulated by thistle-tlssueadditionstothe
success soil.
isas high as grey squirrels'indeclduous The thlstle plants may weaken pasture and, at the
woods.
Captive grey squirrels thrive on a diet of acorns, but sametime, encourage recruitment of theirown
red squirrels have a comparative digestive efficiency klnd. Crowberry (Ernpetrum hevtnaphroditutn) inhibits
of only 57 per cent, seemingly because they are much the growth of Scots pine (Pinus sylvestris) and aspen
less ablethan grey squirrelstoneutralize (Populus trenzula) (Zackrisson and
acorn Nilsson 1992).
polyphenols. A model with slmple competition It releases water-soluble phytotoxic substances from
for
the autumn hazel crop, whlch is eaten by grey squir- secretory glands on the leaf surface. These toxins
rels before the acorn crop, shows that red squirrels interfere with Scots pine and aspen seed germination
are unlikely to persist with grey squirrels in woods on the forest floor.
with more than 14 per cent oak canopy. Oak trees in Aggressivecompetition is not so common as
most British deciduous woods give grey squirrels a is popularly believed. As a rule, organisms avoid
food refuge that red squirrels fail to exploit. Thus, potentlallydangerousencounters.But aggression
red squirrel replacement by grey squmels may arlse does occur. A classic example is two acorn barnacles
solely from feedingcompetltlon,which - Chthamalus stellatus and Balanus halanordes - that
is exacer-
bated by the post-war decline in coppiced hazel. grow intertldal-zone rocks around Scotland. A study
on the shore atMillport(Figure 5.3) showed that,
when free-swimmlng larvae of Chthurnalus decide
Contest competition
to settle, they could attach themselves to rocks down
Contest or interference competition occurs through to mean tide level (Connell 1961). Where Balanus is
directinteraction between indivlduals.Thedirect present,theyattach only down to mean high neap
interactlon may be subtle, as when plants produce tide. During neap tides, the range between low water
toxins or reduce light available tootherplants, or mark and high water mark is at Its narrowest. Young
overtly aggressive, as when anlmal competitors 'lock Balanus grow much faster than Chthumullrs larvae -
horns'. they simply smother them or even prise them off the
Directcompetltion occurs in four duckweed rocks. Thls behaviour is direct, aggressive action to
species (Letnnu mrnor, Lemna natans, Lemna giblla, and secure the available space. When Balunus is removed,
h n n a polyrhiza) (Harper 1761). Lemna minor grows Chthatnalus colonizes theintertidal zone downto
fastest inuncrowded conditions.Under crowded mean tide level. However, wlthout Chthatnalus,
conditlons, when all species are growntogether, Balanus is unable to establish population above mean
Letnnrt ttunor grows the slowest. Nutrlent levels have hlgh neap tlde, seeminglybecause of adverse weather
no effects onthegrowth rates, so thechanges are (warm and calm conditions) promoting desiccation.
caused by competltion for light - the Lemna species
interfere with one another. Mechanisms of coexistence
Several anlmalandplant speciesinterfere with
competitors by chemical means. The chemicals Species coexistence is the
rule in Nature. It
released have an mhibitlng, or allelopathic,impact depends upon avoiding competition and I S achleved
 INTERACTING POPULATIONS 1 17

balanoides Balanus Chthamalus stellatus

A A

Figure 5.3 The vertlcal distributlon of larvae and adult acorn barnacles on intertidal-zone rocks, Millport, Scotland. Two
competing specles are shown - Balanus balanozdes and Chthamalus stellatus. Thais lapillus IS a predatory whelk.
Source: After Connell (1961)

through several mechanisms: the environmentis nor- coexistence of the blackburnlanwarbler (Dmdrozca
mallydiverseand contam hiding placesand food fisca), black-throated green warbler(Dendroicauirens),
patches of various sizes; many species use a range of andbay-breastedwarbler (DmdroicaCastanea). The
resources rather than one; animals have varying food Cape May warbler (Dendrozca tigrznu) is different. Its
preferences and commonly switch diets. The result IS survival depends upon outbreaks of forest insects to
that competitlon 1s often diffuse: a species competes provlde a glut of food. The myrtle warbler(Dendrozca
with many other speclesfor a varietyof resources, and coronata coronata)is notso common as the otherspecies
each resource represents a small portion of the total and IS less specialized.
resource requirements. Some species partltlon resources solely according
to prey size. Among such predatory species as hawks,
larger species normally eat larger prey than smaller
Resource partitioning
species. InNorthAmerlca,thegoshawk (Acczpiter
Speciesavoidnicheoverlap by partitloningthelr gentilis) eatsprey welgh~ng up to 1,500 g (Storer
available
resources
according to
size
and
form, 1966). The smaller Cooper’s hawk (Accrpztw cooperi)
chemical composltlon, and seasonal availability. Five takes prey almostas large, but only very occasionally.
warbler
species(genus Dendroica) live~nspruce It malnlyeatsprey In the 10-200 g range. The
forests in Maine, United States. They each feed In a smaller-stillsharp-shinnedhawk (Acczpzter striatus)
different part of the trees, use somewhat different for- willeatprey up to 100 g, and sometimes a little
aging techniques to find insects among the branches more, but most of Its prey are in the 10-50 g range.
and leaves, and have slightly different
nesting In these specles, the goshawkIS 1.3 tlmes longer than
dates (Figure 5.4) (MacArthur 1958). The differences Cooper’s hawk, which IS itself 1.3 times longer than
In feedingzonesarelargeenough to accountfor the sharp-shinned hawk.
1 18 INTERACTING POPULATIONS

Myrtle warbler Black-throated green warbler Blackburnian warbler

Dendroica coronata coronata Dendroica virens Dendroica fusca

Bay-breasted warbler Cape May warbler

Dendroica castanea Dendroica tigrina

Zones of most intense

feeding activity
B Base of branch
M Middle
T Terminus

6
I
Figure 5.4 Warblers in spruce forests. Maine. Unlted Stares.
Source: After MacArthur (1958)

Character displacement Evolutionary divergence can be inferred only from

the phenomenon of character displacement. This
Competition between two species may be avoided by phenomenonoccurswhena species’appearance or
evolutionary changes in both. Two predators may eat behaviour differs when a competitor is present from
thesame sizeprey andtherefore,allotherfactors when it 1s absent. An example is afforded by the
beingconstant,theyarecompetitors.Underthese beak size of ground finches (Geospizinae) living
circumstances, it would not be uncommon for one on the Gal6pagos Islands (Figure 5.5) (Lack 1947).
species graduallytotackleslightlylargepreyand Abmgdon and Bindloe Islands have three species of
theother species to tackleslightlysmallerprey. Geospzza that partition seed resources according to
The two species would thus diverge by evolutionary size - the large ground finch (Geospzza mgnirostris),
changes. the medium ground finch (Geospiza firm), and the
 119
40

20

40 Albemarle, Indefatigable

40
F
L Chatham Charles,
5 20
:
n.
0
40 1

14 012 10 '16 18 20 22
Beak depth (mm)

Small ground finch

Abingdon 0 I 50 km I Ceospiza fuliginosa

Bindloe 0 0

AAlbemarle

To; Daphne
Medium ground finch
Ceospiza fortis
i

Indefatigable

Large ground finch

mman a dChatham
? Ceospiza rnamirostris

Figure 5.5 Beak m e of ground finches on the Galipagos Islands.

Source: After Lack (1947)
120 INTERACTING POPULATIONS
smallground finch (Geospiza j l l i g t t ~ o ~ u )Gearpizu
. onlylimiting resource. The best competltor is the
rnagnirostris has a large beak adapted to husk large
seeds that smaller finches would fail to break open.
Geusptzu jdigtnusu has a small beak that can husk
smaller seeds more efficiently than the larger species,
Geospzza fh-tts. Geuspizu fortis feeds on Intermediate-
sued seeds. Now, Grusptza rrtugnirostrrs does not reside
onCharles or Chatham Islands. On theseislands,
Geosptza fortis tends to have a heavier beak, on aver-
age, than on Abingdon or Bindloe Islands. O n
DaphneIsland, where Geo.rptzu form lives wlthout
Geosptzu jlliptzosu, Its beak I S intermediate in size
between thetwo specles on Charles andChatham
Islands. O n Crossman Islands, Geospizuj//igtno.ralives
without the other two finches, and its beak is Inter-
mediate in size there. The habitats on all the islands
are very similar, and the explanation for this diver-
gence in beak size 1s thatcompetltlon has caused
character displacement.
Spatial competition
All organisms,andparticularly terrestrial plants
and other sessile species,Interact mainly with their
neighbours. Neighbourhoods may differ considerably
in compositlon, largely because of colonlzation limi-
tation(propagules have not yetarrlved atsome
sites). In areas of apparent uniformlty, thereI S a great
number of ‘botanical ghettos’.This s p t d variety
enablesmore spec~esto coexist than in auniform
environment.Traditionalcompetition theoryholds
that there can be no more consumer species than there
are limiting resources. Thespatialcompetition
hypothesissuggeststhatnelghbourhoodcompetl-
cion, coupledwlthrandom dispersal amongsites,
allows an unlimlted number of specles to survlvr on a
single resource (Tilman 1994). Such rlch coexcstence
occurs because species wlth sufficiently high dispersal
rates perslst in sites that are unoccuped by superior
competitors.
Thespatialcompetltlon hypothesisappears to
explain the coexistence of numerous grassland plant
speciesin theCedar Creek NaturalHistory Area,
Minnesota, United States. The grassland plants com-
pete strongly below ground for nitrogen, whlch is the
littlebluestem (Scbrzuchyrim scopurzurn), a grass.
According to classical competition theory, as there I S
just one limiting resource (soil nitrogen), this species
shouldoutcompete all othersandtake over. In
gardenplots it doesjust that.But in thenatural
grassland it shares theenvironmentwith over a
hundredotherspec~es.The answer tothlsenigma
maylie In the relativeenergy allocation for root
growingand for reproduction. Species investing In
root growth aregood competitors where nitrogen
levels are limiting, but they are also poor dispersers.
Specles investlng in reproductlon, such as the tlckle-
grass (Agro.rti.r . r d r u ) and the quackgrass (Agropyron
repem), though not the best nitrogen competitors, are
gooddispersers. So, the ‘good-dispersers-but-poor-
competitors’ invade abandoned fields immediately,
and are not dominated by the ‘slow-dispersers-but-
good-competltors’untilthlrty or forty years later.
I t is possible,therefore, thatsuperiorcompetltors
are prevented, by thelr poor colonizingabilities,
from occupyingtheentire landscape, andthatthis
provldes sites in which numerous specles of inferior
competitors can persist.
V E G E T A R I A N WILDLIFE:
HERBIVORY
Three forms of populatlon Interaction benefit one
species but are detrimencal to the other. Predation
occurswhen a predatory population(thewinner)
exploits a prey population(the loser). Normally, a
predator has a seriously harmful effect on its prey
- I t kills I t and eats It, but not necessarily in that
order! Thls is classic carnlvory. Plants lose leaves to
herblvores. Herbivory I S thus a form of predation.
Parasitism is a relatively mild form of predatlon,
though i t may still lead to death. Parasitism is like
predatlon but the host (a member of the population
adversely affected), rather than
being
consumed
immediately, is explolted over a perlod.Amen-
salism occurswhen onepopulatlon is adversely
affected by an interactionbuttheotherspec~es is
unaffected.
 INTERACTING
POPULATIONS 121

Plant eaters chavdia) andthewhltemangrove (Azicenna). The

curious New Zealand kakapo (Strigops hahroptifus) is
Types of herbivore a ground-dwellingparrot. It extracts plces from
leaves, twlgs, and young shoots by chewing on them
Herbivores are
animals that
eat live plants. and detaching them from the plant,or it fills its large
Herbivory I S aspeclal kind of predatlonwhere crop wlth browse and retiresto a roost where it chews
plants are the prey. All parts of a plant - flowers, up the plant material, swallows the juices, and ‘spits
fruits, seeds, sap, leaves, buds,galls,herbaceous out’the fibre as dry balls. Members of thegrouse
stems,bark,wood,androots - areeaten by some- family(Tetraonldae)browseonbuds, leaves, and
thing,butnoone herbivoreeats themall.There twlgs of willows and other plants of low nutritive
areseveralspecializedherblvorenlches - grazers, value.
Thelr
digestivesystems housesymbiotic
browsers, grass, grain, or seed feeders (graminivores), bacterla that digest the vegetable materials and are
grainand seed eaters(granlvores), leaf eaters(foli- comparable to the digestlve system of ruminants. A
vores), fruit and berry eaters (fruglvores), nut eaters few birds eat roots and tubers. Pheasants dig Into the
(nucivores), nectar eaters (nectarlvores), pollen eaters soil with their beaks, while others scratch with their
(pollenivores), and root eaters. Detritivores eat dead feet. Some cranes, geese, and ducks are adapted for
plant material and will be discussed in Chapter 6. obtalnlng the roots, rhizomes, and bulbous parts of
A vast army of invertebrate herbivores munches, aquatlc plants.
rasps,sucks,
filters,
chews, andmines Its way Grain,seed,nectar,frult,andnuteatlngare
through the phytosphere. A smaller but potent force specialities ofmany birds. Sometmes, the relationship
of vertebrate herbivores grazes and browses its way between a fruit-producing plant and Its ‘predator’ is
throughthephytosphere(Table5.2).Somemam- close (see Box 5.1). Many mammals eat fruit, butvery
malian herbivore nlches are
unusual. Theglant few arespecialized fruit eaters. An exception is the
panda (Aifuropoda tt/e/anofrf~c.a)I S a purely herblvorous frult-eating bats, whlch includenearly all members of
member of the Carnlvora. Thepalm-nutvulture the Pteropodidae. These anlmals have reduced cheek
(Gypohirmx drzgofetzsrs) I S the only herblvorous teeth as an adaptatlon for their fluld diet.
member of the Falconiformes. It eats nuts of the oil
palm (Elarrs ~ Z ~ I K ~ . and I I . I ) raphla palm(Raphia rzdfu),
Defence mechanisms in plants
as well as fish, molluscs, and crabs. Even Invertebrate
nichescanbesurprising.Anadulttlgerbeetle As prey specles, plants have a distinct disadvantage
(Cil-it/d~la repamid), a common and wldespread water- over the animal herbivore counterparts - they cannot
edge species, was recently seen feedingonfallen move and escape from predators; they have to stand
sassafras frultslyingon a Marylandbeach, United firm and cope with herblvore attackas best they may.
States (Hill andKnlsley1992).This was the first The world is green, so obviously plants can survive
report of frugivory by a tlger beetle, which may be the ravages of plant-eatlnganimals.They do so In
an opportunistic frugivore, espec~ally in the autumn twomain ways. First,someherblvorepopulations
justafteremergence,whenfruitswouldprovlde evolve self-regulating mechanlsms that prevent their
a valuable energy resource before overwintermg. destroylng thelr food supply; or other mechanlsms,
Browsers andgrazers are chiefly mammals, but a partlcularly predation, may hold herblvore numbers
few largereptilesand severalblrdspeciesoccupy In check. Second, all that is green may not be edible
thisniche,too. Some of the geese andgoose-like - plants have evolved a battery of defences against

waterfowlcropgrass.TheSouthAmerlcanhoatzln herbivores.
(O~Z.~~~/J~W hoazin),
/ / U Ja highly
aberrantcuckoo, Plant defences are formldable. Some discourage
malnlyeatsthetoughrubbery leaves,flowers, and herblvores by structuraladaptatlonssuch as thorns
frults of thetall,cane-likewater-arum (Mom+ andspikes.
Some
engage In chemlcal
warfare,
 122 INTERACTINGPOPULATIONS

Table 5.2 Herbivoroustetrapods

Name Order Herbivores

Example Example of
within plant tissue
Order eaten

Amphibians
Anura Frogs,toads Few Commonfrog(Rana Water weed
temporaria) tadpoles
Reptiles
Chelonia Tortoises and turtles Some Giant tortoise Grasses,sedges
(Testudo gigantea)
Squamata Snakes and lizards Few Marine iguana Seaweeds
(Amblyrhynchus
cristatus)
Birds
Anseriformes Ducks and geese Many Southernscreamer Aquatic plants,
(Chauna torquata) grasses, and seeds
Falconiformes Eagles and hawks Few Palm-nut vulture Palmnuts, fish,
(Gypohierax molluscs,crabs
angolensis)
Galliformes Game birds Most Redgrouse (lagopus Heathershoots,insects
lagopus)
Columbiformes Pigeons All Wood pigeon Legumeleaves,seeds
(Columbo palumbus)
Psittaciformes Parrots Most Blue-ond-yellow macaw Fruits.kernels
(Ara ararauna)
Apodiformes Hummingbirds Some Sword-billed Nectar. insects
and swifts hummingbird (Ensifera
ensifera)
Passeriformes Perching birds Many Plantcutter(Phytotoma Fruit,leaves,shoots,
rutila) buds,seeds
Mammals
Marsupialia Kangaroos, etc. Many Honey possum or Nectar, pollen, some
noolbender (Tarsipes insects
spencerae)
Chiroptera Bats Few tong-tongued bat Fruit,nectar,some
(Glossophago insects
soricina)
Dermoptera Flyinglemurs All Flyinglemurs Leaves,buds,flowers,
(Cynocephalus spp.) fruit
Edentata Sloths and Some Threetoed sloths Leaves, fruit
(Xenarthra) armadillos (Brodypus spp.)
Primates Apes,monkeys, Most Human (Homo sopiens) Everything except
andlemurs wood
Rodentia Voles,mice, Most Alpine marmot Grass,plants,roots
squirrels, etc. (Mormota mormota)
 INTERACTINGPOPULATIONS 123

Table 5.2 (continued)

Order Name Herbivores

Example Example of
within plant tissue
Order eaten

Lagomorpha
Rabbits,
hares, hare
Mountain All Fine twigs, sprigs of
and pikas (lepus timidus) bilberry and heather
Carnivora
Dogs,
bears,
cats,
etc. Few panda
Giant Bamboo
(Ailuropoda melanoleuca)
Hyracoidea
Hyraxes
(conies) All hyraxesRock Grass
(Heterohyrax spp.)
Elephants
Proboscidea elephant
African All Grass, leaves, twigs
(loxodonta africana)
ws Sea Sirenia All Manatee Aquatic plants
(Trichechus manatus)
Perissodactyla
Odd-toed
ungulateso All Brazilian tapir Succulent vegetation,
( Tapirus terrestris) fruit
Artiodactyla Even-toed
ungulatesb GoatsAll Everything
(Capra spp.)

Source: Partly after Crawley ( 1 983)

Notes: a Horsesandzebras,tapirs,rhinoceroses
b Pigs, peccaries, hippopotamuses, camels, deer, cows, sheep, goats, antelope

employing by-products of prlmary metabolic path-
ways. Many of these chemical by-products, including
terpenoids, steroids, acetogenins (juglone In walnut
trees), phenylpropanes (in cinnamon and cloves), and
alkaloids (nicotine, morphine,caffeine) are distasteful
or poisonous and are very effective deterrents (see
Larcher 1995: 21).
Herbivores have developed ways of side-stepping
plant defence systems. Some have evolved enzymes to
detoxify plant chemicals. Others time their life cycles
to avoid the noxiouschemicalsin theplants.Two
examples will illustrate these points - cardiac glyco-
sides in milkweed and tannins in oaks.
Poisonous milkweed
Themilkweed Adepzas curassmica is abundant in
Costa Rica andotherparts of Central America. It
contains secondary plantsubstances calledcardiac
glycosides (or cardenolides) that affect the vertebrate
heartbeat and arepoisonous to mammals and birds
(Brower 1969). Cattlewillnot eat themilkweed,
even though it grows abundantly in grass. They are
w s e not to do so, for it causes sickness and occasion-
ally death.However,certain insects, includingthe
larvae of danaid butterflies (Danainae), eat i t without
any deleterious effects. Thedanaidbutterflies are
distasteful to insect-eating birds and serve as models
in several mimicry complexes (p. 111). The danaids
have evolved biochemical mechanisms for eating the
milkweedandstoringthe poisonin their tissues.
Thus,theyacquireprotection from predators from
the plants that they eat.
Unpalatable oak-leaves
The common oak (Quwcus robur) in western Europe is
attacked by the larvae of over 200 species of butter-
flies and moths (Feeny 1970). It protects itself against
this massive assault by usingtannins for chemical
and structural defences. The tannins lock protelns In
complexes that mects cannot digest and utilize, and
they also make leaves toughandunpalatable.The
herbworous attackers partly get round these defences
124 INTERACTING POPULATIONS

Box 5.1
THE H A W T H O R N A N D THE fruitssimply fall to the ground. Robins dropped
A M E R I C A N ROBIN: 20 per cent of thefruitsthattheypicked.They
n FRUIT-FRUGIVORE SYSTEM defaecated or regurgitated 40 per cent of the swal-
lowed fruits (seeds) beneath parent bushes. Bushes
European hawthorn (Crataegus monogyna) grows in with more fruit were visited more frequently, had
western Oregon, United States. Only one frugivore, a greaterdispersal success (more seeds weredis-
the American robin (Turdus migratwirts), forages on persed),and had seedsdispersedmoreefficiently
the fruits, making this an unusually straightforward (a greater proportion of seedswasdispersed, and
Fruit-frugivore system (Sallabanks 1992). The fruits the seeds weremore successfulin propagating).
carry seeds thataredispersed by theAmerican Theoptimalfruitingstrategy for theEuropean
robin,who bears them away fromtheparent hawthorn is, therefore, to growas big as possible as
bush. Dispersal efficiency is low. An average 21 per quickly as possible by delaying fruiting until later
cent of seeds are dispersed each year. Most of the in life.

by concentratlng feeding in early spnng, when the
leaves areyoungand, becausethey contain less
tannin, less tough. They also alter their life cycles
in summer and autumn - many late-feeding insects
overwinter as larvae and complete their development
on the soft and tasty springleaves. Some insect species
do feed on oak leaves in summer and autumn. These
tend to growvery slowly, which may be an adaptation
to a low-nitrogen diet when protelns are lockedaway
in older leaves.
Herbivore and plant interactions
Herbwores interact with plants In two chlef ways.
First, in non-interactive herbivore-plant systems,
herbivores do notaffect vegetationgrowth, even
when there are very large numbers of them. Second,
in interactive herbivore-plant systems, herbivores
affectvegetationgrowth.Twoexampleswillillus-
tratethesedifferentsystems - seedpredationand
grazlng.
Seed predation
Herblvores that feed upon plantseeds (seed predators)
do not normally influence plant growth. Birds eatmg
Grazing
plant seeds have no effect on plant growth, though
they could influencethe long-term survivalof a plant In interactive herbivore-plant systems, the herbivores
populatlon by eating a large portlonof the annualseed affect the plants, as well as the plants affecting the
production. On the other hand, plants do influence
herbivorepopulatlons - seed predatorpopulatlons
will be low In tlmes of seed shortages. Britishfinches
feed elther on herb seeds or on tree seeds. The herb-
seed feeders have stable populatlons, whereas the tree-
seed feeders have fluctuatingpopulatlons(Newton
1972).Herbsareusuallyconsistentintheannual
number of seeds produced, but tree-seed production
is variable and,In some cases, irregular. Variable tree-
seed production IS illustrated by the North American
piiion pine(Pznus edulis).The piiion pine yleldsheavy
a
cone crop at irregular intervals. The crop satiates its
Invertebrate seed and cone predators. It also allows
efficient
foraging by piiion
Jays (Gymnorhmus
cyanocephalus),which dispersethe seeds and store them
In sites sultablefor germination and seedling growth
(Ligon 1978). Many other trees are mast fruiters. In
mast fruiting, all trees of the same specles wlthin a
large area produce a big seed crop in one year. They
then wait a long time before producing anything other
than a few seeds. The advantage of such synchronous
seeding is that seed predatorsaresatlatedandare
unlikely to eat the entire crop before germlnatlon has
begun.
 INTERACTING POPULATIONS 125

herblvores. As arule,plants have moreImpacton is thesmallestcarnlvore ever - andthe‘largeand

herbivore populatlons than herblvores have on plant equally deadly’ - the African lion (Panthem (eo). Some
populations - plant abundance, quality, and distribu- fossil carnivores wereawesome In slze and possibly
tiongreatly affect herbivorenumbers.Nonetheless, inspeed.Themostfamousexamples, even before
herbivory can affect plant populations. This is the case their appearance in Jurassic Park, are the dinosaurs
where Vegetation is grazed. Sheep, for example, are Tyrannosuwl~srex and Velociraptor. Butmorerecent
‘woolly lawnmowers’ thatmalntalnpasture.The avian predators were just as terrifying. The phorus-
effectsof grazlng on vegetation are plalnest to see rhacoids were a group of large, flightless, flesh-eating
wherea fence separatesgrazedand ungrued areas. blrds (L. G. Marshall 1994). They lived from 6 2
The grazed area consists malnly of grasses, while the milliontoabout 2.5 million years ago in South
ungrazed area I S domlnated by shrubs and tall herbs. America and became the dominant carnivores on that
Given enough tlme, the ungrazed area would revert continent. They ranged In helght from 1 to 3 m, and
to woodlancl. Domestic livestock sometimes has such wereable to killandeatanimals the size of small
a largeinfluenceonplantcommunlttesthatthe horses.
herblvoremaybeldentlfiedfrom the vegetation it
produces (Crawley 1983: 295). In the United King-
Carnivorous specialization
dom, dock (Kwttex obtusifolius) is abundant In horse
pastures; silverweed (Potentilh ansenna) IS common Allanimals,exceptthoseatthe very top offood
where geese graze. Spiny shrubs and aromatlc, stlcky chains, are eaten by a predator. The zoological menu
herbs flourish wheregoatsgraze In Mediterraneanis vast, butpredatorstend to specialize.Carnivore
maquls. nlchesIncludegeneralflesh-eaters(faunavores), fish
Patterns ofherblvore-plant Interaction are complex eaters(piscivores),blood
feeders (haematophages
andgeneralizatlon I S difficult.Populationmodels, or sanguivores), egg eaters(ovavores),Insecteaters
similar to those used to study predator-prey relatlon- (insectlvores), ant eaters(myremovores),andcoral
shlps, suggest some basic possibilities (Box 5.2). eaters (corallivores). Omnivoresare
unspeclalized
carnivores-cum-herbivores. Scavengers feed ondead
animal remains. Coprophages feed on dung or faeces.
HUNTERANDHUNTED:CARNIVORY A few carnlvorousnichesarehlghlyspecialized
andratherunusual.Thereare few sanguivores,the
most notorlous (apart from Count Dracula) are the
Flesh eaters
New World vampire bats. The sharp-beaked ground
Carnlvores are animals that eat anlmals. Carnivory I S finch (Geosp~zad;ffili.r), one of the Galipagosfinches,
predationwhereanimalsaretheprey.Carnlvory obtains some its nourlshment by bltlng the bases of
Includespredatorseatingherblvores as well as top growlng feathers on boobies ( M a sp.) and eatlng the
predators eating predators. A predator benefits from blood that oozes from thewound.Coprophagy IS

interactlon wlth aprey - I t gets a meal; the prey does morewidespreadthanmightbesupposed. Several
not benefit from interactlon with a predator - it nor- blrds eat animaldung.The Ivory gull (Papphila
mally loses Its life. Rareexceptlonsarelizardsthat e b l m e a ) feeds on the dung dumped on ice by polar
lose thelrtails to predatorsbutotherwlsesurvive bears, walruses, and seals. Puffins andpetrelseat
attacks. Predation is plam to see and easy to study. It whale dung floatlngonthewater. Certain vultures
is evident when one anlmal eats another, often in a andkltes feed solely onhumanandcanine faeces
gory spectacle. around natlve villages. Myrmecophagy (ant eating) is
Predatorscome in a varietyofshapesandsizes. found in several specles in Australia, South America,
Highlights from a predator catalogue might be the Africa, and Eurasia that are adapted to feed upon ants
‘small but deadly’ - the least weasel (Mustela mtwli.r) and termms (Figure 5.7). Extremeadaptatlons for
126 INTERACTING
POPULATIONS

Box 5.2
POPULATION MODELS OF In these
equations, P is the
plant
population, H is
HERBIVORE-PLANT SYSTEMS population, K is the carrying
the herbivore capacity
of plants, and a , 6 , c, and dare parameters: a is the

prey systems.The plantis the prey and the herbivore herbivores; b is the depression of the plant pop- '
is the predator. Theequationsdevisedtostudy lationperencounterwith a herbivore,ameasure
predators and their prey (p. 130) maybe adapted to of herbivore searching efficiency;c is the increase in
cnrrl.rherhirrnrec nrevrino .-.n nlancr ThP pnllatinnr the herhivnre nonulacinn ner encounter withdants.

mpulationi are (Ciwley 1983: 249): the decline in herbivores (death rate) in the absence
of plants.
dt'fdt = [aP(K - P)]IK - bHP The parameters and
the
carrying
capacity
affect system stability and steady-state population
dHldt = cHP - dH. numbers. In brief, increasing the intrinsic growth
rate of plants, a , increases systemstabilityand

(a) a-0.2 1 a-0.4

b-0.001
1 b-D.005

k ,,""""""""

1
c=o.001

2,400 - (d K-500 K-1,000

1,200 -

lai. . . . 1

Figure 3.G Herbivore-plant interactions.

Source: After Crawley (1983)
 INTERACTING
POPULATIONS 127

herbivoresteady-statedensity,buth&no effect abundanceand so reducessystemstability, but

3n plant abundance (Figure 5.6a). Increasing herbi- increasessteady-stateherbivorenumbers(Figurt
Bore searching
efficiency, b, reduces,somewhat 5.6~).Increasing the carrying capacityofthe environ.
wrprisingly, herbivore steady-state density, but has ment for plants, K, increases herbivore steady
no effect onsystemstability or onsteady-state statedensityand reduces systemstability,but
plant abundance (Figure 5.6b). Increasing herbivore paradoxically, has noeffectonsteady-stateplant
growth efficiency, c, reduces steady-stateplant abundance (Figure 5.6d).

Figure 5.7 Ant-eating mammals fromfourorders:Xenarthra - all members of the family Myrmecophagidae, which
includes the giant or great anteater (Myrmecopbugutndactylu);Tubulidentata - the aardvark (Otycteropus ufer);Pholidota -
eight species of pangolin (Mums);and Monotremata - the Australian and New Gumean spiny anteaters or echidnas,
Tucbyghus urnledus and Zuglossus bruijnnr. The numbat (Myrmecobrurfu~ciutuJ), which IS not illustrated, is an Australian
marsupial anteater.
Source: Animal drawings from Rodriguez de la Fuente (1975)

this diet areseen in teeth, jaws, and skulls. The teeth
are reduced to flat, crushing surfaces, as in the aard-
vark (Oyc-teropus afer), or they are missmg, as in the
giantorgreatanteater (Myrmecophaga trihtyla).
Tongues are long, mobile, and worm-like, wlth un-
common protrusibility (‘stick-out-ability’). Enlarged
salivary glands produce a thick, sticky secretion that
coats the tongue. Ant-eating mammals also tend to
have elongatedsnoutsandpowerfulforelimbs for
diggmg and rlppmg open nests.
Some plants are carnivorous, including the great
sundew (Drosera anglica) and the greater bladderwort
(Utrimlaria vulgarzs). Pitcherplants of thegenus
Heliamphora (Sarracenraceae) in the Guayana High-
lands of Venezuela have carnivorous traits gaffe et
al. 1992). Heliamphora tatez IS atruecarnwore.It
128 INTERACTING POPULATIONS

attracts prey through speclalvlsual andchemlcal primates, that are eaten by the tlger (Punfhera tqr/.r),
signals,trapsandkillsprey,digestspreythrough the leopard (Pnnthwu pardm), andthedhole (Cmm
secreted enzymes, contalns commensal organlsms, and ulpznrrs) (KaranthandSunquist1995).Thethree
has wax scales thataid prey captureandnutrient predatorsshowsignificantselectivityamong prey
absorptlon.Other speclesof Hem~u7nphoru possess species. Gaur (Bos ga:ccNrrds) I S preferred by tlgers,
all thesetraltsexceptthey seem nelther to secrete whereas wild pig (Sns .rcrofu) I S underrepresentedin
proteolytlc enzymes nor to produce wax scales. The leopard diet, and the langur or leaf monkey (Pre.rhyrr.r
patternofthecarnlvoroussyndromeamong Heliu- entellrrs) I S underrepresented In dholediet.Tigers
tnpboru species suggests that the carnlvorous hablthas selected prey weighingmorethan 176 kg, whereas
evolved in nutrient-poorhabitats to ~mprovethe leopard and dhole focused on prey in the 30-175 kg
absorption of nutrlents by capturingmainlyants. slze class. Average weights of prlncipal prey killed by
Only Hefiatnphoru tutrr further evolved mechanlsms tiger, leopard, and dhole were, respectlvely, 91.5 kg,
for rapidassimilatlon of organicnutrientsand for 37.6kgand 43.4 kg.Tigerpredation was biased
capturlng a variety of flying Insects. The carnivorous towards adult males in chital o r axis deer (Axis axr.0,
tralts are lost in low light conditions, lndicatlng that sambar ( C m m /tnrtafor), andwild pig, andtowards
nutrlent supply I S limltlng only under fast growth. young gaur. Dholes selectively preyed on adult male
chltal, whereas leopards did not. If there ischoice,
large carnlvores selectively kill large prey, and non-
Prey switching
selective predatlon patterns
reported
fromother
The switchlngof preyfollows the principleof optlmal tropical forests may be the result of scarcity of large
foraglng.Predators select the preythatprovides prey. Because availability of prey In the appropriate
the largest net energy gain, considering the energy S I Z C classes
is not a limiting resource, selective
expendedincapturingandconsumingprey.This predatlon may facilirate large carnlvorecoexlstence In
usLrally means the preythathappened to be most Nagarahole.
abundant at the time. It alsomeans that predators
normallytake very young, very old,orphysically
Carnivore communities
wrakenedprey.Somecarnlvores show local feeding
specializationswlthintheirgeographicalrange.In Carnlvore communitles normally consistof predators
Mediterraneanenvlronments,theEuropeanbadger with overlapplng tastes. For this reason, they posses
(hiehtnefes), a carnivore species wlth morphological. complex feeding relationships. A dry tropical forest
physlologlcal, and behavioural tralts proper to a feed- in the2,575-km’ Hual
Kha
KhaengWildlife
Inggeneralist, prefers to eattheEuropeanrabbit Sanctuary, Thailand, contained twenty-one carnivore
r h ) et a/. 1995). It will eat specles from five families(RablnowitzandWalker
( O v y d u p l s c ~ / n i ~ ~(Martln
otherpreyaccording to thelravailabilitywhenthe 1991). The carnivores fed onat least thirty-four
rabbit klttens are not abundant. The badger I S a poor mammalspec~es, as well as birds,lizards,snakes,
hunter, and its ability to catch relatlvely Immobile crabs, fish, insects, andfrults.Nearlyhalf the prey
baby rabbits may explaln the curlous speclalization. identlfiedin
large
carnlvore faeces, which was
mainly deposlted by Aslatlc leopard (Panthcva puvdus)
andclouded leopard (Neofifis nehrrlo.ru), was barking
Prey selection
deer (Mztntracr4.r mrtntjak), with sambar deer (Cert~rts
Where several carnlvores competefor the same range rrnrcdor), macaques (Mucaca spp.), wild boar (Sus
of prey specles, coexistence I S possible if each carni- scrofu), crestless Himalayan porcupine (Hyrtrix
vore selects a different prey. In the tropical forests of hodpni), andhogbadger (An.tonyyx collurrs) being
Nagarahole, southern India, there I S a wlde range of important secondary prey items.Murld rodents,
large mammalian prey specles, mainly ungulates and especially the yellow raph-rat (hfuxotnys mr+r), and
 INTERACTING
POPULATIONS 129

the bay bamboo-rat (Cut/non/yshadim), accounted for density. The pars include: sparrow-hawk (Europe),
one-third of identified food items in small carnivore muskrat-mink(centralNorthAmerica), hare-lynx
faeces. Non-mammalpreyaccounted for ~ u s tover (boreal North America),muledeer-mountamlion
one-fifth, and fruit seeds for one-eighth, of all food (Rocky Mountains), white-tailed deer-wolf (Ontarlo,
items found In small carnlvore faeces. Canada), moose-wolf (Isle Royale), caribou-wolf
(Alaska), and white sheepwolf (Alaska).
The ratlonalebehlnd predator-prey cycles is
Predators and prey deceptivelystraightforward.Consider a population
Commonly,predatorandpreypopulatlons evolve of weasels, the predator, and a population of voles,
together.Predatorswhicharebetterable to find, the prey.Ifthe vole populationshouldbelarge,
capture,andeatpreyare morelikely to survive. then,withaglut of food scamperingaround,the
Natural selection tends to favour those hunters’ traits weasel populatlon will flourish and increase. As the
withinpreypopulations.Inturn,predationpushes weasel population grows, so the vole population will
thepreypopulation to evolve ina directlonthat shrlnk. Once vole numbers have fallen low enough,
favours individualsgood at hldingandescaplng. the preyshortagewill cause a reductlon Inweasel
Overthousandsof generations, evolutionary forces numbers. The vole population, enjoylng the dearth
actinguponpredatorand prey populatlons lead to of predators, will then rlse agaln. And so the cycle
elaborateandsophisticatedadaptations.These fea- contlnues.
tures are seen In the social hunting behavlour of lions Althoughthe
rationalebehind predator-prey
and wolves, thefoldingfangsandvenom-injectlng cyclesis plausible, sustalned oscillations in predator
apparatus of viperlnesnakes, ; m i spidersandthelr and prey systems are not always the result of popula-
webs. tlonInteractions.Canadianlynx (Lynx crraadntms)
populatlons show a nine to ten-year peak In denslty
(Figure 5.8). This cycle isrevealedby lynxpelt
Predator-prey cycles
records kept by Hudson’s Bay Company in Canada.
Thereare severalpairs of predatorsandpreythat The Canadian lynx feeds matnly upon the snowshoe
appeartodisplay cyclicalvariationsinpopulatlon hare (Lep4.r tirrrrrdmrrs), which also follows a ten-year

n - Snowshoe hare (Lepuscanadensis)

Canadian lynx (Lynx canadensis)

1850 1860 1890

1880
1870 1900 1910 1920 1930
Year

F&re 5.8 Cycles In thepopulartondenslty of theCanadian lynx (Lynx ~-mrrr/rn.rrj)and the snowshoe hare (Ltpi.r
c-nnnr/ens/.r),as seen in the number of pelts received by the Hudson’s Bay Company
S o / m e : After MacLulich ( I 937)
130 I N T E R A C T I N GP O P U L A T I O N S

cycle. Field investlgationssuggested

that
the cycle 25
snowshoe
in hare density is correlated with the 20
availability of food (theterminaltwigs of shrubs
and trees). The lynx populatmn, therefore, follows l5
3 2-
swings insnowshoe hare numbers,and does notdrive 2 10

them. 2%
5
Oscillations in predator-prey systems were
modelled in the 1920s (Lotka 1925; Volterra 1926, 0
19.31). Refined versions of these
produced three Important
models
findings.
First,
populat~oncycles occuronly whenthe prey'scarry-
have
stable
-
5c
111

2
lo:

8
m
0 Sprlng density
Autumn denslv
P Predator

Ing capaclty (inthe presence of predators) IS large. $:

- V
-
Thisfinding goes againstintuition.It I S called the 2': 4 -

'paradox of enrichment'andstatesthat an increase

in carrylng capaclty decreases stability. Second, the
-$' 2 -

population cycles are stable only when the prey 0 I I

1992 1987 1982 1977 1972
populatlongrows faster thanthepredatorpopula-
tion, or when thepredators arerelatively Inefficient F ; , ~ ~ ,j.9
,.~ ~ l , cycles
I,reclarors
~ ~ ~ ~ ~ I,rey, d ~ l ~ , ~ k ~
atcatchlngthelr prey. Thlrd,some predator-prey westernFinland. The prey I S voles (/Lfimtmspp.) nncl the
displaythe features Of chaotic dynamics. predator 1s the least weasel (AINI/E/Nt / / l , d / / ~ ) .
S r m w : Afrer Hanski et a / . (199.3)

Chaos in Finnish weasel and vole populations

scopic protozoans, Puruttzmwt cut~dut/~ttt and Didinium
Long-termsrudies of predator (weasel) and prey nuwttm. Dzdinrt~ttr prey voraciously on Pavuttzrl.zrm.
(vole) dynamics In Finland have revealed populatlon Inoneexperiment, Puvutttecztm and Didinium were
cycles of three to five years (Figure 5.9). A predator- placed In a test-tube containlng a culture of bacteria
prey
model with seasonal effects Included (by supported In a clear homogeneous oat medium. The
allowlng weasels to breed only when the vole density bacteria are food for the Puvuttreciz~nr.The outcomewas
exceeds a threshold)predictedpopulatlon changes that Didinzum ate all the Purutnnrc.izrtt/ and then died
that closely resembled the observed changes In of starvation.This resultoccurredno matter how
weasel and vole populations (Hanskl et ul. 1993). The large the culture vessel was, and no matter how few
study suggested rhat the cycles In vole populations Didiniutn were introduced. In a second experiment, a
are drlven by weasel predation, and are chaotlc. These little sediment was added to the test-tube. This made
findings are supported by further workon boreal the
medium heterogeneous and afforded some
voles and weasels that shows theimportance of refuges for the prey. The outcome of this experiment
multispecies predator-preyassemblages with field- was the extinction of the DIdinzutt/ and the growthof
vole-type rodents as the keystone species (Hanskl and the prey populatlon to its carrying capacity. In a third
Korpimaki 1995; Hanskl and Henttonnen 1996). experiment, the prey and predator populations were
'topped up' every three days by adding onePnrumeciuttz
and one Ddirrirrttz to the test-tube. In other words,
Geographical effects
immigratlon was Included In theexperlment.The
outcome was two completecycles of predator andprey
Laboratory experiments
populatlons.
. .
Gauseconcludedthat,wlthoutsome
Gause (1934) studied a simple predator-prey system form of externalinterferencesuch as immigration,
in the laboratory (Figure 5. IO). He used two micro- predator-prey systems are self-annihilating.
 INTERACTING POPULATIONS 131

0- Paramecium caudaturn
0""- 0 Didiniurn nasutum

1 immigration 'waves'

6 9 12 15 18
Time (days)

F i p r e 5.10 Outcomes of Gause's laboratory experlments wlth the protozoans Purumeum carrdufutu (prey) and D/dir/ttm
nu.r/mm (predator). (a) Oatmedium wlthout sediment.(b) Oat medium with sediment. ( c ) Oat medium wlthout sediment
but wlth immigratlons.
Source: After Gause ( 1 934)

Later,laboratoryexperimentsusing anorange thedistancebetweenorangesslmplylengthened

+ herbivorous mlte + carnivorous mite food cham
showed that coexistence was possible (Huffaker
1958). Whentheenvironment was homogeneous
(oranges placed close together and spaced evenly), the
predatory mite (Typhlodromus ocudentalis) overate its
phytophagousprey (Eotetvanyhrrssexmalxlatuss) and
died out, as Gause's Dtdinzum had done. Increasing
the time to extinction.Butwhentheenvironment
was made heterogeneous, the
systemstabilized.
Severaldifferent'environments'wereconstructed.
The basic'landscape' was fortyoranges placed on
rectangular trays like egg cartons, with some of the
orangespartly covered with paraffin or paper to
limlt the available feeding area. Complications were
1 32 I N T E R A C T I N GPOPULATIONS

1,200~ - Herbivorous mite (Eotetranychus

sexrnaculatus) t

0 0
0 10 20 30 40 50 60
Time (weeks)

F&re 5. I I Population cycles in a laboratoryexperlmenc where a predatory mite, Typhlodrourtr~ o&dmtali.<, feeds
upon another mite, Eotetrunyrhtrr sextuuctrluttrs. The environment is heterogeneous, conslsclng of 252 oranges wlrh one-
twentleth of each orange available to the prey for feeding.
Sotrrce: After Huffaker et a/. (1963)

introduced by using varying numbers of rubber balls thepopulationsmaintainedsustamedoscillatlons

as 'substitute' oranges. In somecases, enure new trays
oforangeswereaddedandartificialbarriers of
Vaseline constructed that the mites could not cross.
Stabilityoccurredina252-orangelandscapewith
complexVaselinebarriers. The preywereable to
colonizeoranges In ahop,skip,andjumpfashlon,
and to keep one step ahead of the predator, whlch
exterminated each littlecolonyofprey it found.
During the seventy weeks before the predators died
outandtheexperlmentstopped,threecomplete
predator-prey population cycles occurred (Figure
5.11).
Nearly forty years after Gause's original work, new
experlmentsusing Paratt/eL.rtrm artrelict as preyand
Dir/in/uttI as predator managed to stabilize the system
(Luckinbill 1973, 1974). When Paran/ecr/ttt/ was
grown wlth Didit/z/m in a 6 ml of standard cerophyl
medium, Didil/r/tm ate all the preyina few hours.
Whenthemedium was thickenedwithmethyl
cellulose to slow down the movements of predator
and prey alike, the populations went through two or
threedivergingoscillationslasting several
days
before becommg extinct. When a half-strength cero-
for thlrty-threedays before theexperlment was
concluded. A mathematicalmodelofthesystem
suggests that the stability arlses because the Increase
in the cost-benefit ratioofenergyspentsearching
to energy gained capturlng prey apparently inhiblts
the predator searching at low prey densities (G. W .
Harrison1995).
An ambltlous setof laboratory experiments studied
competltlon and predation at the same time (Utlda
1957). The system contained a beetle, the azuki bean
weevil (Callosohrttdws cbrnensts), as prey that was
provldedwithanunlimlted food supply.Butthis
coleopteranparadise had a 'sting in the tail' In the
form of two competing species of predatory wasp -
Neowrolucut.\ t ~ ~ ~ t t ~ e z o p band
u p sHeterospiluj prosoprdzj.
The wasp specles had slmilar life histories and
depended upon the beetleas a food source. Durlng the
four years oftheexperlment,whichrepresented
seventy generatlons, all three populatlons fluctuated
wildly but managed to coexist (Figure5.12).The
wasp population Huctuatmns were out of phase wlth
one another. Thls was because Heterospilrts was more
efficient at finding and exploltlng the beetle when It

phylmedium was thickenedwithmethylcellulose, was at low densitles,while Neocatolacws was more

 INTERACTING POPULATIONS 133

- Azukibeanweevil (Callosobruchus chmensrs)

.-c
v1
8oo
600
1 Parasitic wasp (Neocato/accus mamezophagus)
...-..A....-.
---D-- Parasitic wasp (Heterospilusprosopidis)

C
U
.- 400
Y
-
2
n
0
a
200

0
0 10 20 30 40 50 60
Generation

Figure 5 . I2 Populatlonchanges in a laboratoryexperlmenr wlth a beetle host, the azuki bean weevil (Cullosobrrrrbru
and two parasitic wasps, Neorutolarctrs tt~att~ezophagrrs
C/J~NK?ISIS), and Heterospihs prosnprdis. The experiment ran for four years.
All three populatlons fluctuated considerably, bur they did suTvIve.
Sorrrre: After Utida (1957)

efficient athigher prey densities.Thecompetitive
edge thus shiftedbetween the two wasp specles as the
beetle density changed wlth tlme (owing to density-
dependentchangesinreproductionrateandthe
effects of the two wasp populations). The stability of
the system was thus thepurely result of predatory and
competltlve biotic interactions.
rates between landscape patches. They also reveal the
curious fact that some metapopulations may perslst
withonly'smk'populations, in whichpopulation
growth is negative In the absence of mlgration.
However, long-term persistencerequires some local
populationsbecoming large occaslonally (Hanski et
dl. 1996a).

Mathematical 'experiments'
Intheory,geography IS a crucialfactor In the co-
existence of predator and prey populations. A simple
mathematicalmodel showed that when predators
and prey interact within a landscape, coexistence is
rather easily attalned fJ. M. Smith 1974: 72-83). It
is favoured by prey with a hlgh migration capacity,
by cover or refuge for the prey, by predator mlgration
during a restrictedperiod, and by a large number
of landscape'cells'. Furthermore, if thepredator's
mlgratlon ability is too low, I t will become extlnct.
If thepredator'smlgrationability is hlgh, co-
existence is possible if the prey is equally mobile.
Simplemathematicalmodels of predator-prey
interactlons in a landscape have evolved into sophls-
ticated metapopulatlon models. These, too, stress the
vltal role of refuges for prey species and mlgratlon
L I F E A G A I N S T LIFE:ALTERNATIVES
TOCHEMICALCONTROL
Pests are organlsmsthat interfere withhuman
actlvitles,and especially with agriculture. They are
unwelcome competitors, parasites, or predators. The
chief agricultural pests are insects (that feed mainly
on the leaves and stems of plants), nematodes (small
worms that live mainly In the soil, feeding on roots
and other plant tissues), bacterlal and vlral diseases,
andvertebrates(mainlyrodentsandblrds feeding
on gram and frult). Weeds are a malor problem for
potentlal crop loss. A typical fieldis Infested by ten
to fifty weed specles that complete with the crop for
light, water, and nutrients.
Pest control I S used to reduce pest damage, but
even with the welght of modern technology behlnd
134 INTERACTING POPULATIONS

it, pest control is not enormously successful. In the used toregulatepest populations. Two approaches
Unlted States, one-third of the potential harvest and exist - mundative blocontrol and classical biocontrol
one-tenth of the harvestedcrop I S lost t o pests.A (Harris 1993). In inundative control, an organlsm I S
controloperatlon 1s successfillif the pestdoesnot applied In the manner ofaherbicide. Like a herbicicle,
cause excessive damage.It I S a failure if excessive the control agent IS usually marketed by Industry. In
damage I S caused. Just how much damage is tolerable classical biocontrol, an organlsm ( o r possibly a
dependsontheenterprlseandthepest.An insect virus) I S established from another reglon. The pest is
that destroys 5 per cent o f a pear crop may be Insig- keptin check inclefinitely,usually by government
nlficant in ecological terms, but I t may be disastrous agencles actlng in the public Interest. Both biocontrol
for a farmer's margin of profit. On the other hand. apprwaches, in the rlght clrcumstances, are effective
a forest Insect may strlp vast areas of trees of thelr means of pest control and brlngfew harmful envlron-
leaves, but the lumber Industry will not go bankrupt. mental Impacts, as thefollowing
exmnpleswill
Pestsarecontrolled In severaldifferent ways. Theshow.
blanketapplicatlon of toxlc chemicals calledpesti-
cides has lnimlcalslde-effects ontheenvlronment.
Prickly-pear cactus in Australia
Several otheroptions for pestcontrolareavailable
(Table 5.3). The prlckly pear (Op/t)/tm.rtr~to),a cactus, I S natlve to
North and South Amerm. It was brought t o eastern
Australia In 1839 as a hedgeplant.It spread fast,
Biological control
formlng dense stands, some1-2 m high and too thlck
Biological pest control pits predator specles against for anybodytowalkthrough. By 1900, I t Infested
prey species - parasites, predators, and pathogens are 4 million ha. Control by poisoning the cac'tus wits not

Table 5.3 Main pest control techniques

Control method Intermediate

r-strategist
K-strategist
pests
pests pests

Pesticides
(use chemical
Early widespread Selective
pesticides
Precisely
targeted
compounds
to kill pests application
applications
based on
directly) monitoring on forecasting
Biological (use natural enemies, Introduction or
viruses, bacteria, or fungi) enhancement of natural
enemies
Cultural
(change
agricultural
Timing,
cultivation, + t Changes in
or otherpracticestoalterthe and rotation agronomic practice,
pest's habitat) destruction of alternative
hosts
Resistance (breed animal General, polygenic + t Specificmonogenic
cropplant
and varieties
resistance resistance
resistant to pests)
Genetic (sterilize pest Sterile mating techniques
population to reduce its growth
rate)

Source: After Conway ( 1 98 1 )


economically feasible - clearing infestedland with
poison was far more costly than the worthof the land.
Searches in natlve habltats of the prlckly pear were
begun in 1912 to find a possiblebiological control
agent. Eventually one was found -a moth, Cactoblastrs
cactorum, native to northern Argentina, the larvae of
whlch burrow into the pads of thecactus,causing
physical damage and introducingbacterial and fungal
infections. Between 1930 and 1931, when the moth
population had become enormous, the prlckly pear
stands were ravaged. By 1940, the prickly pear was
still found here and there, but in very few places was
I t a pest.
A different controlagent, Dac.tylopirrs
oprmtrae,
was used in an area of New South Wales, Australia,
where Cactoblastzs cactorurn had not been not a success-
ful biological controlagent(Hosking et al. 1994).
Dactyloprrls oprrntiae reduced a prickly pear (Optrntra
strrcta var. strrcta) populatlon In the central tablelands
of New South Wales.
False ragweed in Australia and India
The false ragweed (Parthenirrm hysterophorrrs),a native
of Central Amerlca, IS a problem weed of Australian
rangeland,particularly In Queensland. Following
field surveys in Mexico, therustfungus Pr~c.inra
abrrrpta var. partheniicola was selected as a potential
biological control agent (Parker et al. 1994). One]so-
late was chosen for furtherinvestigation. Infection
with the rust hastened leaf senescence, significantly
decreased the life-span and dry weight of false rag-
weed plants, and led to a tenfold reduction in flower
production.Subsequentstudies showed thatthe
rust was sufficlently host-speclfic to be consldered
for introduction. In Bangalore, India, Cassra rmrjora,
a leguminous undershrub of some economicvalue,
has replaced, over a five-year period, more than 90
per cent of the false ragweed on a 4,800m’ site Uoshi
1991). Leachates from Cas.ria rrm/lora are allelopathic,
inhibiting false-ragweed seed germination and ham-
perlng the establishment of a summer false-ragweed
generation. The colonies of Ccrssra unzjora are robust
enoughtoprevent a false-ragweed generation from
forming below them in wlnter.
INTERACTINGPOPULATIONS 135
Pests in the Mediterranean
The whitefly, Parabemisia nlyricae, is one of the most
serious citrus pestsin the eastern Mediterranean
region of Turkey. In 1986, a host-specific parasitoid
of Parabenzisranryricae, the
aphelinid Eretnrocerrrs
debachr, was imported fromCalifornia(Sengonca et
al. 1993). In the following years Parabenzisia rnyrrcae
populations were rapidly reduced In all citrus
orchards where the parasitoids were released. Eretnru-
cerus debachi was a good disperser,well adapted to the
climatic conditlons. Since its successful colonization,
the whitefly IS no longer a serious pest.
In Cyprus, the black scale (Sai-rsetza oleae) IS a pest
primarily of olive tree (Olea europaea).I t attacks several
otherplants,includingcitrus trees and oleander
(Nermn oleander). Two parasitoids (Metaphycrts bartlettr
and Metapbyeus helvolus) were Imported, mass-reared,
and permanentlyestablished in the island (Orphanides
1993).Following limited releases of these parasltolds,
black scale populations fell from outbreak levels to
almost non-existence. Black scale populations have
stayed low since parasltold releases were discontlnued.
Genetic control
Anotheralternatlveto chemical control of pests is
genetic control. One method of genetic control is
to breed sterileorganisms. For example, if gypsy
moth (Lymantria dispar) pupae are irradiated with a
sterilizing dose of gamma radiation and mated with
normal females, the first generation male moths are
sterile (Schwalbe et a/. 1991). However,difficulties
associated with large-scale deployment of partially
sterile males renders genetic control impractical on a
large scale.
Genetlc control is also achieved by breeding crop
plantsthat aremoreresistant to pests. It was this
control technique that, in 1891, came to the rescue
of the Frenchwine industry. Phylloxera is an aphid
natlve to Amerlca. It lives In galls on leaves and roots
of vineswhere i t cannot be reached by sprays. I t
multiplies prodiglously. Vines infected by Phylloxera
become stunted and die. In 1861, it was accldentally
136 INTERACTING POPULATIONS
introducedinto Languedoc,France. Two decades
later, four-fifths of the Languedoc vineyards had been
devastatedand every wine-growlng area of France
was infected;noremedyhad been found;andthe
outlook was bleak for wine drinkers. In 1891, it was
discovered that the American vine(Vitzs lahrusca)was
almost immune to Phylloxera. Sclons of the European
vme (Vitis vinifwa) were grafted onto American root
stocks to produce a hybrid vine that, if not entirely
immune, was affected far less seriously.
Integrated pest management
Modern pest control involves integrated pest
management. This I S an ecological approach to pest
management that brings together at least four tech-
niques. First, I t uses natural pest enemies, Including
parasites, diseases, competitors,andpredators(bio-
logical control). Second, I t advocates the planting of
a greater diversity of crops to lessen the possibility
that a pest will find a host. Third, it advocates no
or little ploughing so that natural enemies of some
pests have a chance to build up in the soil. Fourth,
i t allows the applicatlon of a set of highly specific
chemicals, used sparmgly and judiclously (unlike the
old applicatlon method that tended to be profligate).
Integrated pest management involves the use of
chemicals, the development of genetically resistant
stock, blologlcal control,and land culture. Land
culture is the physical management of the land -
whether and how it is ploughed, what kind of crop
rotation used, the dates of plantmg, and basic means
of handling crop harvests to reduce presence of pest
in residues and products sold.
Integrated pest management has been used to
tackle theorientalfruitmoth (Grapholithamolesta),
which attacks several frult crops (Barfield and Stimac
1980). Themoth is prey toa species of braconid
wasp, Ma(.rocentrus anr)liwous. Theintroductlonof
the wasp into fields and orchards helped to control
themothpopulation.But an interesting discovery
was made: the efficacy of the wasp in peach orchards
was increased when strawberry fields lay nearby. The
strawberry fields are an alternativehabltat for the
wasp and help it to overwinter.
SUMMARY
No population can exist isolation - it needs to inter-
act with others. Population interactions take several
forms but fall into two categories - co-operation and
competitlon. Co-operation occurs to varying degrees.
Weak co-operation (protocooperatlon) is beneficial to
both populations, but it is not obligatory - the inter-
actmg species would survive without one another. It
includesmimlcry.Mutualism is an extreme form
of protocooperation in which theinteraction is
obligatory.Commensalism occurswhen one of the
co-operating species benefits andthe
other is
unharmed by the assoclation. Competitlon occurs
where species interactions are detrimental to at least
one of theinteractants.Thecompetitive exclusion
prlnciplestatesthat no two species may occupy
exactly the same nlche. If they should occupy similar
nlches, cornpetition willoccur. Competition takes
the form of scramble competition (for resources) or
contest competitlon (sometimes involving aggressive
action between individuals). Coexistence of compet-
Ingspecies is achieved by resource partitionrng,
characterdisplacement,andspatialheterogeneity.
Herbivory I S the predatlon of plants by plant-eaters.
The plant kingdom has evolved defences to foil thetr
herbivorous adversaries. The defences include struc-
turaldeterrentsand chemicalarsenals. Herbivores
interactwithplants In several characteristic ways.
This is seen in patterns of seed predationand
grazing. Carnivory is the predatlon of one anlmal by
another. There are several carnlvorous specializations,
some rather blzarre. Evolutron has finely tuned carni-
vore communitiestopromote coexistence through
prey switchlngand preyselection. Predatorsand
their prey sometimes display cycles In population
numbers. Such cycles aresometimestheresult of
food availability, though chaotic behaviourin some
predator-preysystems does appearto rest wlthln
theinteractingpopulations themselves. Stability in
predator-preysystems is more readily attained in
heterogeneous environments. Life may be pitted
against life in anattempttocontrolagricultural
pests. Biological control is a useful alternativeto
pesticideapplication.
Genetlccontrol is another
 INTERACTING
POPULATIONS 137

option. Biological control I S usually applied withm a FURTHER READING

broader system of integrated pest management.
Crawley, M. J.(1983) Herbivory: The Dynarnm oj
Animal-Plant lnteractions (StudiesinEcology,Vol.
EQSUSEASYT I O N S lo), Oxford: Blackwell
Sclentlfic Publications.

1 What are theresultsofinterspecific Grover, J. P.(1997) Resource Competition, New York:

competition?
2 What are the advantages and
disadvantages of speciesco-operation?
3 What are the pros and consof
biological control?
Chapman & Hall.
Jackson, A. R. W. and Jackson, J. M. (1996)
Environmental Science: The Natural Environment and
Hutnan Impact, Harlow: Longman.
Kingsland, S. E. (1985) Modeling Nature: Episodes rn
the History of Population Biology, Chicago and London:
University of Chicago Press.

Kormondy, E. J. (1996) Concepts OfErology, 4th edn,

Englewood Cliffs, NJ: Prentice Hall.

MacDonald,D. (1992) The Velzjet Claul:ANatural

History of the Carntzjores, London: BBC Books.
 6

COMMUNITIES
Communities consist of several interacting populations. Ecosystems are communities together
w i t h the physical environment that sustains them. This chapter covers:

the nature of communities and ecosystems

ecological roles in communities
feedingrelationships
biologicaldiversity

SOCIAL A N D PHYSICAL chalk(Figure 6.1). A variety of soils have formed

CONNECTIONS: COMMUNITIES uponthese rocks. The soils support manytrees
AND ECOSYSTEMS (Figure 6.2). The main ones are oak (chiefly Q/rwcrrs
robrdr), hornbeam (Carpinus betrhs), and silverblrch
An ecosystem is a space in which organisms interact ( B e t d a penddu). The herb layer is varied, the main
withoneanotherandwiththe physical envlron- communitiescorrespondingtothe chief habitats
ment. A communityis the assemblage of interacting - the forest floor, streamsandotherdamp places,
organismswithinan ecosystem. Communities are paths, clearings, the woodland edge, a marshy area,
sometimes called biocoenoses, with each part given and achalky area. It consists of flowering plants, ferns
a separate name - phytocoenose (plants), zoocoenose and horsetails, mosses and liverworts, andfungi.
(animals), micro-biocoenose (micro-organisms) There are 283 plant species, at least 3 liverwort and
(Sukachev and Dylis 1964: 27). Where the physical 12 moss species, and 302 fungi species. A few lichens
environment supporting the community is included, grow on tree trunks and buildings.
the term biogeocoenose is used (and is a vowel-rich The plants and fungi support a great diversity of
and somewhat awkward alternatlve to the term eco- insects and woodlice. There are predators(ground
system). Ecosystems and communities rangein size beetles, burying beetles, ladybirds, and
others),
from a cubic centimetres to the entire world. defoliators (leaf beetles and weevils), seed borers (wee-
vils),barkborers(weevils and barkbeetles), wood
borers ('longhorn' beetles, 'ambrosia' beetles, and click
A local ecosystem beetles), root feeders (weevils and click beetle larvae
The Northaw Great Wood, Hertfordshire, England, -wireworms), and woodlice. There are butterflies and
is adeciduous wood, with a small plantation of moths, snails and slugs. There are two amphibians -
conifers. I t occupies 217 ha In the headwaters of the the common toad (BNfO hdo) and the common frog
Cuffley Brook dralnage basin. The eastwards-draining ( R u m temporurza); and three reptiles - the warty newt
streams have cut into a PebbleGravel plateau that sits (Trzturmcristatm), the slow-worm (Angzrzs /ragili.r),
at around 400 m. They have eroded valleys in London and the rlngedo r grass snake (Natrix natrm). There are
Clay, Reading Beds (sands), and, at thelowest pomts, fifty specles of breeding blrds and some twenty-two
 COMMUNITIES 139

.....
..._
.....
.."

! 6.1 Northaw Great Wood, Henfordshlre, England. (a) Geology. (b) Soils.
: (a) After Sage (1966b). (b)After D. W King (1966)
140 COMMUNITIES

Figure 6.2 Tree distribution In the Northaw Great Wood. Hertfordshlre, England.
Source: After Horsley (1966)

visitors. Insummer,thedommantspeciesinthe Most ecosystems are heterogeneous and involve a

breeding communlty are chaffinch (Fringiffacoefebs), mosaic of individuals, populations, and habitats. The
willow warbler (Phyffoscopus trochilus),blue tlt (Parus Northaw Great Wood 1s mainly an oak-hornbeam

caerufus), great tlt (Parus major), robin (Eritbams woodland, with stands of silver birch on more acldic
rubecufa),blackbird (Turdus merula),wren (Trogfodytes and better drained soils. However,it contains patches
troglodytes),dunnock or hedge sparrow(Pruneffamoa'u- and corridors. The corridors are formedby paths and
faris), blackcap (Syfvza atrzcapiffa),gardenwarbler the streams. The patches include cleared areas and
(Syfvzaborzn), chiffchaff (Phyffoscopuscoffybzta),greater areas where other tree species dominate. There are,
spottedwoodpecker (Dendroropos m j w ) , nuthatch for example, three stands of ash (Fraxznus excelsior),
(Sitta europaea), redstart (Phoenicurus pboenznrrus),and three stands of beech (Fagus syfvatzca), three stands of
tree plpit(Antbus trzvzafis).Twenty-three wild spec~es aspen (Populus tremufa), and seven stands of sweet
of mammal are recorded (Table 6.1). Six mammalian chestnut (Castanea sativa).
orders are represented - Insectivora (shrews, hedge- Withinthe wood, several communlties exist.
hogs, and voles), Chiroptera (pipisrrelle and noctule A good example is the bracken (Pteridium aquilinzum),
bats), Lagomorpha(haresandrabbm),Rodentia wood anemone (Anemone nemwosa), and
bluebell
(squirrels,rats,mice,andvoles),Carnivora (foxes, (Hyarzntbozdes non-srrzpta) community (Sage 1966a:
badgers, stoats, and weasels), and Artiodactyla (fallow 14). These species manage to live together as a plant
deer and muntlac deer). association by dividing the habitat to avoid undue
 COMMUNITIES 141

Table 6. I Mammals in Northaw Great Wood, Hertfordshire, England (to 1966)

Order Family Species

lnsectivora Erinaceidae
Hedgehog (Erinaceus europaeus)
europaea)
(Talpa
Mole
Talpidae
Soricidae
Common shrew (Sorex
araneus),
pygmy shrew
(Sorex minutus), water shrew (Neomys fodiens)
Chiroptera Vespertilionidae
Common
pipistrelle (Pipistrellus pipistrellus), noctule
bat (Nyctalus noctula)
Lagomorpha Leporidae
Rabbit (Oryctolagus cuniculus), brown hare
(lepus
europaeus)
Rodentia Sciuridae Grey squirrel (Sciurus carolinensis)
Bank vole (Clethrionomys glareolus), field vole
(Microtus ogrestis), water vole (Arvicola terrestris)
Wood mouse (Apodemus agrestis), yellow-necked
mouse (Apodemus flavicollis), brown rat (Rattus
norvegicus)
Dormouse (Muscardinus avellanarius)
Canidae
Carnivora Red fox (Vulpes vulpes)
Stoat (Mustela erminea), weasel (Mustela nivalis),
badger (Meles meles)
Artiodactyla
Cervidae
Fallow deer (Dama dama), Reeve's muntjac
(Muntiacus reeves4

competition for resources. Brackenrhizomes may ments.IntheNorthawGreatWood,tree,shrub,

penetrate about 60 cm. The bluebell bulbs are not
nearly so deep. The wood anemone taps the top 6 cm
of soil. The wood anemone is a pre-vernal flowerer
(March to May), whereas thebluebell is a vernal
flowerer (April to May). Both complete assimilatlon
before the bracken canopy develops In June, so avoid-
ing competition for light. The bluebell is the only
species that can withstandbracken,but if bracken
growth IS especially vigorous, the accumulations of
fallen fronds exclude even the bluebell.
Communities possess vertlcal layers. Thls stratifi-
cation results primarily from differences in light
intensity. In somecommunities,distlnct layers of
leaves are evident - canopy, understorey, shrub layer,
and ground layer. Animal speclesaresorted among
the layers according to thelr food and cover require-
field, and ground layers are present. Each layer acts
as a kind of sernl-transparent blanket and modifies
theconditions below. Light levels arereduced, as
is precqxtationintensity, from the canopy tothe
ground.
The horizontal distributlon of species wlthln
communlties is more complicated.Twoextreme
sltuatlons exist. First, a gradient is a gradual change
in a species assemblage across an area. In the Northaw
Great Wood, the domlnant treespecles change along
a toposequence. Silver blrch on summlts gives way to
oak and hornbeam In mldslope posltmns and along
the valley floor. Second, a patch I S a clustering of
spec~es Into somewhat distinctlve groups. Although
tree communities do gradually alter along envlron-
mental gradients In the Northaw Great Wood, there
142 COMMUNITIES

are also stands of trees that form distinctpatches autotrophs produce organic materials by oxidation
(Figure 6.2). Gradients and patches are two ends of of inorganic compoundsand
do not requlre
a continuum and the sub~ect of a lively debate in sunlight.
vegetation sclence. 2 Consumers (heterotrophs). These are organisms
thatobtalntheir food andthus energy from
the tissues of otherorganisms,eitherplants or
The global ecosystem
anlmals or both - herblvores, carnivores, and top
All livlng thmgs form the biosphere. The blosphere carnivores. Consumers are dividedInto several
Interacts with non-livlng things in Its surroundings trophlc levels. Consumersthat eat livingplants
(air,water, soils, andsediments)to win materials are primary consumers or herblvores. Consumers
and energy. The ~nteraction creates the ecosphere, of herbivores are the flesh-eating
secondary
whlch is defined as life plus life-support systems. It consumers or carnivores. In some ecosystems,
consists of ecosystems - Individuals, populations, or there are carnivore-eating carnivores;theseare
c o m m u n ~ t ~ einteractlng
s wlth thelr physical environ-tertiary consumers or top carnivores.
ment. Indeed, the ecosphere is the global ecosystem 3 Decomposers(saprophages)anddetritivores
(Huggett 1995: 8-11; 1997). (saprovores). Decomposers dissolve organlc
Communltiesand ecosystems possess pcopertles matter,whiledetritivores break it intosmaller
thatemerge from the indiv~dua~organisms.These pieces and partly digest it.
emergent properties cannotbe measured in individ-
uals - they are community properties. Four important Autotrophs produce organic m a t e r d , t h e consumers
communityproperties arebiodiversity, production eat It, and the decomposers and detritivores clean up
andconsumption,nutrient cycles, and food webs. the mess (excrement and organlc remains).
Thesecommunitypropertiesenablethe biosphere
to perform three important tasks (Stoltz et ai. 1989).
Producer society: community
First, the blosphere harnesses energy to power itself
production
andbuildup reserves of organicmaterial. Second,
it garners elements essential to life from the atmos-
Primary producers
phere, hydrosphere, and pedosphere. Third, I t is able
to respond to cosmic,geological, and biological Greenplants use solar energy, in conjunctionwith
perturbatlons by adjusting or reconstructing food carbon dioxide, minerals, and water, to build organlc
webs. matter. The organic matterso manufactured contains
chemicalenergy. Photosynthesis IS the process by
whlch radiant energy I S converted Into chemical
PRODUCING AND CONSUMING: energy.Production I S thetotal biomassproduced
COMMUNITY ROLES by photosynthesis withln a community. Part of the
photosynthet~cprocess requires light, so it occurs
All organisms have a community and an ecologlcal only duringdaylighthours.Atnlght,the stored
role. Some organismsproduceorganiccompounds, energy I S consumed by slow oxdation,a process
and some break them down. The chlef roles are as called respiration in Individuals and consumption
follows: in a community,
Sunlight comes from above, so ecosystems tend to
I Producers (autotrophs). TheseInclude all photo- have a vertical structure (Figure 6.3). Theupper
autotrophic organisms:green plants,eukaryotic production zone is rich in oxygen. The lower con-
algae,blue-greenalgae, andpurpleandgreen sumption zone, especially thatpart in the soil, I S
sulphur bacterta. In some ecosystems, chemo- rich incarbon dioxlde.Oxygen is deficient In the
 COMMUNITIES 143

Ecospheric Biogeochemical Spatial structure

zones cycles

Gaseous storage zone

Air Atmosphere

Production zone
Photosynthetic plants dioxide,
Carbon
dominant
Compensation level
Regeneration zone
Animals and
microconsumers
dominant
Solid storage zone
Soils and sediments
Circulatory mode
Ecosystem
. . . . . .I. . 4 ...............
....................................................................
...................
r )Production 7 .........................
:.i. .~ ~ d i ~ ~ i j ; ~ ~ r ~ i i , i l i i i i i i j i : i j
... ........
water, and
nutrients
Lithosphere
-L
Consumption .J
Circulation by roots,
movements, migration of stems, gravity, and
organisms,and (in reefs moving air
and algal mats) diffusion
Aquatic: Terrestrial:
the hydrobiosphere the geobiosphere
(lakes, rivers, oceans, (forests, grasslands,
deserts, and tundra)
.
"

Sake: From Huggett (1997) partly aker Odum (1971)

consumption zone, and may be absent. Such gases as is similar In temperate forests, though the absolute
hydrogen sulphlde, ammonla, methane, and hydro- valuesarelower.Grassland plantsareall leaf and
genareliberatedwherereducedchemlcalstates root - there is negligible branch and trunk. Tundra
prevail. The boundary between the productlon zone and semi-desert plants largely made of green plant
and the consumption zone, whlch is known as the blomass and contain little supporting tmue.
compensation level, lies at the point where there
isjustenoughlight for plantstobalanceorganlc
Primary production
matter production against organlc matter utilization.
Phytomass IS the living material In producers. It The greenplantsthatformtheproduction zone,
normally excludes dead plant materlal, such as tree becausethey producethelrown food fromsolar
bark, dead supportlng tlssue, and dead branches and energy
and
raw
materlals,
are
called photo-
roots. Where dead bits of plants are included, the autotrophs. The amountof organlc matter that they
term standing crop IS applied.Evergreentreesin syntheslze per unit time is gross primary produc-
tropical forests are largely made of dead supportlng tion.Most of thismatter IS created In theplant
structures (branches, trunks, roots); about 2 per cent leaves. Some of I t is transported through the phloem
is green plant blomass (leaves) (Figure 6.4). The rel- to other partsof plants, and especially to the roots, to
ative proportionof leaves, branches, trunks, and roots drlve metabolic and growth processes.
 144 COMMUNITIES

Leaves
Branches
Stems
Rook

and‘Temperate’
Tundra
‘Tropical’
Grassland
forest forest semidesert

”
and’Temperate’
Tundra
‘Tropical’
Grassland
esert forest forest
Figure 6.4 The distribution of biomass In different biomes. (a)Total carbon (glgatonnes, Gg) stored In leaves, branches,
stems, and roots. (b) Carbon stored per untt area (kglm’) in leaves, branches, stems, and roots. 'Tropical forest’ tncludes
tropical forest, forest plantations, shrub-dominated savannah, and chaparral. ‘Temperate forest’ lncludes temperate for-
est, boreal forest, and woodland.
Source: After data In Goudriaan and Ketner (1984)

Net primary production is the grossprlmary kilometre ofice(assummgthedenslty IS 1 g/cm3)

production less the chemical energy burnt In all the wouldwetgh 1 billion
tonnes (1 petagramme).
actlvltles that constitute plant resplratlon. Net pn- A block of Ice 1 km hlgh and restlng on an area of
mary production IS usually about 80 to 90 per cent 15 km x 15 km would have about the same mass as
of gross prlmary production. The mean net global the global net prlmary productlon.
primacy productlon 1s 440 glm2/yr (Vitousek et al. In terrestrlal ecosystems, the mam producers are
1986), or about 224 petagrammes (1 Pg = 224 X greenplants(megaphytes),with the lowerplants
loLsgrammes = 224 billion tonnes) (Figure 6.5). It playmg a mlnor role. Land plants produce on average
IS difficult to comprehend such vast figures.A cublc 899 glm’lyr (Vitousek et a/.1986). That IS a total of
 COMMUNITIES 145

2'oool

I.
E
v
1,500
C
.-0
Y

500
m
1

FiEure 6.5 Mean net prlmary. production

_ for the biosphere and Its component blomes.
So&: Data from Vitousek et al. (1986)

about 132 billion tonnes for the entlre land area. In
aquatic ecosystems, the main producers are autotro-
phic algae. These unicellular and colonial organisms
are suspended in the water and are partof the plank-
ton. Aquatic plants produce on average 225 g/m2/yr
(Vitousek et af. 1986). That is a total of 92.4 billion
tonnes for theentlrewater area. Ofthlstotal,
91.6 billion tonnes are produced by marme plants,
and a mere 0.8 billion tonnes by freshwater plants.
A very tiny part of global net primary production
is contributed by chemolithotrophlc bacteria using
chemlcal reactions around vents In the sea floor or in
soils.
Theworldpattern of terrestrial
netprimary
productionisshowninFigure 6.6. I t mirrors the
simultaneous availability of heatand
molsture.
Production is hrgh in the troplcs, warm temperate
zone, and typlcal temperate zone, and low In the arid
subtropics, continental temperate regions, and polar
zones.
Consumer society: community
consumption
Consumers
The materialproducedbyphotosynthesisserves as
a bas~clarder for an entire ecosystem - it is used as
anenergy-richreserve of organicsubstancesand
nutrients that IS transferred through the rest of the
system. The potential chemical energy of net primary
production is available to organlsms that eat plants,
both livmg plant tissue and dead plant tissue, and
indirectly therefore to anlmals (and the few plants)
that eat other animals. All these organisms depending
upon other organlsmsfor their food are heterotrophs
or consumers. They are browsers, grazers, predators,
or scavengers. They
include
microscopicorgan-
isms,such as protozoans,andlargeforms,such as
vertebrates. Themajorityarechemoheterotrophs,
but a few speclalizedphotosyntheticbacteriaare
146 COMMUNITIES
\
g d.rn/rn2/year
I
2,500
2,000
Figure 6.6 The world pattern of terrestrial net primacy productlon. Units are grammes of dry matter per square metre
per year (g d.m/m*/year).
Soutw: From Huggett (1997) after Box and Meenterneyer (1991)
photoheterotrophs.ThestoredchemicalenergyIn
consumers is called secondary production. There are
two broad groupsof consumers. Macroconsumers or
biophages eat living plant tissues.Microconsumers
or saprophages slowly decompose and disintegrate
the waste products and dead organic matter of the
biosphere.
The availablephytomass In water is small,but
the primary productlon (the amount of dry organic
matter produced In a year) is relatively large because
aquatlc plants multlply fast. Animals eat the plants
and incorporate much of the primary production in
their bodies. For thls reason, thesecondary produc-
tion or zoomass (animalbiomass) In aquatic eco-
systems IS commonly more than fifteen tlmes larger
thanthephytomass.Thesituation is differentin
terrestrial ecosystems. Muchof the phytomass consists
of non-photosynthetlc tissue, such as buds, roots, and
trunks. For thls reason, the standing phytomass is
always very large, especially in woodlands. Primary
production, on the other hand, is relatively small -a
modest amount of dry organic matter is created each
year. A mere 1 per cent, or thereabouts, of the phyto-
mass IS eaten by anlmals living above the ground. The
zoomass is therefore small, being about 1 to 0.1 per
cent of the phytomass.Biomass IS the weightor mass
of living tissue in an ecosystem - phytomass plus
zoomass. It has anenergycontent,whichmay be
thought of as bloenergy.
The human harvest of plants for food, fuel, and
shelteraccounts for about 4.5 per cent of global
terrestrlal net production (Table 6.2). Land used in
agrlculture or converted to other land uses accounts
for about 32 per cent of terrestrial net primary pro-
duction. All human activitieshave reduced terrestrial
netprlmaryproduction by around 45 percent.
This probably amounts to the largest ever diversion
of primaryproductiontosupportasingle species
(Vitousek et al. 1986).
Ecosystems also contain ‘geomass” soils (including
 COMMUNITIES 147

Table 6.2 Human appropriation of net primary production in the 1980s

Manner of consumption production

primary Net

Total mass Percenta e of

(billion tonnesj 3
terrestria net
primary production
~ ~ _ _ _ _ _ -~_
Used by humans
Plants eaten 0.8 0.48
Plants fed to domestic animals 2.2 1.67
Fish eaten by humans and domestic animals 1.2 -
Wood For paper and timber 1.2 0.9 1
Fuel wood 1 .o 0.76
I
To to 7.2 4.54"
Used or diverted
Cropland 15.0 1 1.35
Converted pastures 9.8 7.42
Other (cities, deforested) 17.8 1 3.48
Total 42.6 32.25
Used, diverted, or reduced
Used or diverted 42.6 32.25
Reduced by conversion 17.5 1 3.25
Total 60.1 45.50

Source: After Diamond (1987)

Note: Excluding fish

litter and dead organic matter), sediments, an, and
water that harbour a supply of water and nutrients
(macronutrients,micronutrients,
andother trace
elements).
Decomposers a n d detritivores
Saprophages include decomposers (or saprophytes)
anddetritlvores (or saprovores). Decomposers are
organismsthat feed ondead organlcmatterand
waste products of an ecosystem. They do so by secret-
Ing enzymes todigestorganlcmatter In theirsur-
roundings,andsoaklng u p the dissolved products.
They are mainly aeroblc and anaeroblc bacterla, pro-
tozoa, andfungl. An example is the shelf fungus
(Truruetes zersrdor). This grows on rotting trees and is
Important In decomposlng wood. A very few ani-
mals,such as tapeworms,andsome green plants,
such as theIndianpipe, also obtaintheir food by
diffusion from the outside.
Detritivores are organisms thatfeed upon detritus.
They Include beetles, centipedes, earthworms, nerna-
todes, and woodlice. They are all mlcroconsumers.
Detritivores assist the breakdown of organic matter.
By chewmg and grinding dead organlc matter before
ingestion, they comminute I t and render it more
digestible. When egested, thecarbon-nltrogen (C/N)
ratio I S a little lower, and the acldity (pH) a little
higher, than In the ingested food. These changesmean
that the faeces provlde a better substrate for renewed
decomposer (and notably bacterial) growth. Succes-
sively smallerfragments of dead organicmatter
are passed through successwely smaller detrltivores,
after havmg been subjectto decomposer attackat
each stage. The resultI S a commlnutlon sp~ral (Figure
6.7).
148 COMMUNITIES

-
r

1- Herbivory
I -
I Autotrophs I t I
b
- Herbivores I

i
c Mycorrhiza

I/ SaprovoreS
tr

1 I' Decomposers

Saprophytes
I-
t t Decomposition,
humification cycles
Nutrient

Uptake
""" """"""""
I

++
uptake by
saprophytes

Mineralization
I
Saprophagy
I
(Comminution spiral)

-.
I
I

organic 1
i Deadmatter I
I
I

I
- 1

2
I I
Decomposition I I Turnover

1 Deadanimalsandfaeces

2 Leaf fall, timber fall, root sloughage,

dead plants, wash-out, pollen, seeds

Figure 6.7 Grazlng and detrltal feeding relatlons In an ecosystem, showing commmutlon splrals.
Source: After Huggett (1980)

Humificatlon accompanies comminution and de-
composition to produce a groupof organic compounds
called humus. Humus is in a sense the final stage of
organic decomposmon, but it is also the product of
mlcroblalsynthesisand IS always subjecttoslow
mlcroblal decomposition, the end productof whlch IS
stable humus charcoal. Humus decomposltion in the
temperate zones is Incomplete and humus is brown
or black. Decomposmon is usually more advanced In
the tropicsandhumus may becolourless.Mineral
nutrients are released into the soil solution during
decomposltion, a process styledmineralization.
 COMMUNITIES 149

Recycling
machines:
ecosystem thrlr
grandest
At scale,
blogeochemical cycles
turnover exogenlcAn Earth.
involve
entire the Involving
cycle,
thetransportandtransformation o f materlals near
Biogeochemicals the
surface,
Earth’s isdistlngulshed
normally from
a slowerand less well understoodendogeniccycle
The biosphere IS madeofthree main elements- hydro-
Involvingthe lower crustandmantle. Cycles of
gen (49.8 per cent by weight), oxygen(24.9 per cent),
carbon, hydrogen, oxygen, and nitrogen are gaseous
andcarbon ( 2 4 . 9 percent). Several otherelements
cycles - their component chemicalspecles are gaseous
arefoundinthebiosphere,andsomeofthemare
for a leg of the cycle. Other chemical species follow
essential to biological processes - nitrogen (0.27 per
asedimentary cycle because they d o notreadily
cent), calcium (0.07.3 per cent), potasslum (0.046 per
volatilizeandareexchangedbetween the blosphere
cent),silicon (0.033 percent),magneslum (0.031
and its environmentin solution.
per cent), phosphorus( 0 . 0 3 per cent), sulphur(0.017
Mineralscyclethroughecosystems,thedrlving
percent),andalumlnium (0.016 percent).These
force beingthe flowof energy. T h e c ~ r c ~ ~ l a tof ~on
elements, except aluminium,are the basic ingredients
mlneral elements through ecosystems Involves three
for organlc compounds, around which blochemistry
stages - uptake,turnover,anddecompositlon. Sol-
revolves. Carbon,hydrogen,nltrogen,oxygen,sul-
utes and gases are taken u p by green plants, the rate
phur, and phosphorus are needed to build nucleic acids
of uptake broadly matching biomass production rate,
(RNAandDNA),amino acids(protelns),carbo-
and incorporated into phytomass. Oxygen is released
hydrates (sugars, starches, and cellulose), and liplds
In photosynthesis. The remalnlng mlnerals are either
(fatsandfat-likematerials). Calcl~lm,magnesium,
passed on to consumers or else returnedto soil
andpotassiumarerequired In moderateamounts.
and water bodies when plants die or blts fall off. The
Chemicalelementsrequiredinmoderateandlarge
mlneralsintheconsumerseventuallyreturn to the
quantities are macronutrients. Morethanadozen
soil, sea, or atmosphere. Figure 6.8 summarlzes some
elementsarerequlredintraceamounts,including
major mineral cycles.
chlorine, chromium,copper,cobalt,iodine,iron,
The distribution of blogeochemicals wlthin eco-
manganese, molybdenum, nickel, selenlum, sodium,
systems varies among bmmes. Figure 6.9 shows the
vanadium, andzlnc.
These are micronutrients.
amount of carbon stored In biomass, litter, humus,
Functional nutrientsplay some role In the metabolism
and stable humuscharcoal In the majorecozones. This
ofplantsbut seemnot to be ~ndispensable.Other
glvesagoodindication of how organlcmatter I S
mineralelements cycle through liv~ng systems but
apportioned in different ecosystems. Biomass carbon
have no known metabolic role. The blosphere has to
obtainallmacronutrlentsandmicronutrlentsfrom is twenty to forty times greaterIn forests than in other
its surroundings. ecosystems, Humus carbon I S hlghest In temperate
forests and grasslands. It I S lower in tropical forests
because I t I S rapldly decomposed in the ycar-round,
Biogeochemical cycles hotandhumidconditions.Stable-h~lmus-charc~al
There is in the ecosphereaconstantturnover of carbon, whlch degrades exceedingly slowly,I S hlghest
chemicals. The motive force behlndthesechemical in troplcal forests, and is still slgnllicant in temperate
cycles is life. In addition, on geological time-scales, forests and grasslands.
the cyclesare Influenced byforces in the geosphere
producingandconsuming rocks. Biogeochemical
cycles, as they are called, involve the storage andflux
of all terrestrial elements and compounds except the
inert ones. Material exchanges between life and life-
support systems are a part of b~ogeochem~cal cycles.
I'

EATERS A N D THE E A T E N F
: OOD
C H A I NA
S N FDO O W
D EBS
Food webs
An ecosystem contains two chief types Of food webs:
agrazing food web(plants, herbivores,carnivores,
top carnivores) and a decomposer or detrital food web
(Figure 6.7).
Grazing food chains and webs
This simple feeding sequence
plant + herbivore + carnivore + top carnivore
IS a grazing food chain. An example is
leaf + caterpillar + bird + weasel.
Usually though, food and feeding relations in an eco-
system are more complex because there I S commonly
a wlde variety of plants available, different herbivores
prefer different plant species, and carnivoresare
likewlseselectlve about whichherblvoretheywill
consume. Complications also arise because some
animals,the omnlvores,eat bothplantandanimal
tissues. For all these reasons, the energy flow through
an ecosystem is inmost cases better described as a
food web.
Figure 6.10 shows the food web for Wytham
Wood,Oxfordshire,England.TheIncoming solar
energy supports a variety of trees, shrubs, and herbs.
Oak (Qlrevc.us spp.) IS the dominant plant. The plants
are eaten by a variety of herblvores. Several inver-
tebrates feed on plant material, and especially leaves.
Thewintermoth (Operophteru brrmatu) and the
pea-green oak twist (Tortrixvzridanu) are examples.
The herblvoresare prey to carnivores, including
spiders, parasites, beetles, birds, and small mammals.
Cyzenrs ulbimns I S a tachlnld fly and a specific parasite
of thewintermoth.There are several predatory
beetles in the litter layer, the commonest large ones
belng Philonrhlrs decorm, Feronia nladida, Felonia mlu-
nuriu, and Ahax purullelopipedlrs. Titmice feed partly
on plants(e.g. beech mast), partlyoninsects, and
partly on spiders.The main specles are the g r a t
COMMUNITIES 151
tit (Parzrs tnalor) and
the
blue tit (Partis cawuleus).
Small
mammals Include bank
the vole (Clethrionomys
glureolus) and the wood mouse (Apodonzrs sylvatzc~rs).
Top carnivores include parasites and hyperparasltes,
shrews,moles, weasels, andowls.Thecommon
shrew (Sorex maneus), pygmy shrew (Sorex nunutus),
and mole (Tulpa enropaeu) dominate the ranks of top
carnivore. They are all eaten by the tawny owl (Strzx
a h o ).
Decomposer food chains and webs
Dead organic matterand
other waste products
generallylie upon and wlthln the soil. As they are
decomposed,minerals are slowly released that are
reused by plants. There are complex food and feeding
relations among the decomposers and a decomposer
or detrital food chain is recognmd, the organisms
in which are all microconsumers(decomposers and
detrltlvores).
O n land, litter and soil organic matter support a
detrital food web. In Wytham Wood, decomposers
(mainly bacteria and fungi) slowly digest the litter.
Litterfragments are eaten by detritivores - earth-
worms, soil insects,and mites. Predatorbeetles eat the
detritivores, thus linking the grazlng and detrital
food
webs. Inaquatlc ecosystems, the producers (malnly
phytoplankton) live in the upper illuminated areas of
water bodies. They are eaten by anlmal microplankton
andmacroplankton. In turn,theanimalplankton
are eaten by fishes, aquatic mammals, and birds that
take thelr prey out of water. All the organlsms In the
detrltal aquatlc food cham are eventually decomposed
in the water and in sediments on lake, river, or sea
bottoms.
Ecological pyramids
Energy and biomass are distributed among producers,
herblvores, predators, top predators, and
decomposers.
Their distributions bothresemble a pyramid -energy
and blomass become less in movlng from producer, to
herbivore, to carnivore, totop carnlvore. This I S
because at each higher trophic level, there is SLICCCS-
slvely less energy and blomassavailable to eat. In
152 COMMUNITIES

6oma.s

-In Litter
Humus
Stable humus charcoal

‘Troplcal‘ ‘Temperate‘
Grassland Tundra and Agncultural Human
forest forest semldesett land use land use

Stable humus charcoal

‘Troplcal’ ‘Temperate‘
Grassland Tundra and Agricultural Human
forest forest semidesert land use land use

Figure 6.9 Carbon stored In biomass, litter, humus, and charcoal in the major ecozones. (a) Total carbon (gigatonnes,
Gg).(b) Carbonper unit area (kg/m2). ‘Tropicalforest’ includes troplcalforest,forest plantations, shrub-dominated
savannah, and chaparral. ‘Temperate forest’ includes temperate forest, boreal forest, and woodland.
Source: After data In Goudrlaan and Ketner (1984)

CedarBogLake,Minnesota,thenetproduction distribution of biomass - narrow at the bottom and

figures (in kcal/m2/yr) are: producers 879, herbivores
104,and carnivores 1 3 (Lindemann1942).The
biomass of each trophic level could be measured in
the field, or could be derived by convertlngthe
production figures to biomass. Each gramme of dry
organic matter is equivalent to about4.5 kcal, so each
kilocalorie unit is equivalent to about 0.222 kg. The
b~omassesfor Cedar Lake Bog (in kg/m2) are there-
fore approximately:producers195,herbivores23,
and carnivores 3. This conversion to biomass does not
change the shapeof the pyramid.
Someecosystems have an Inverted
pyramidal
wide at the top. In the English Channel, there are
4 g of phytoplankton per m2 and 21 g of zooplank-
ton per m2. Clear-water aquatic ecosystems have a
lozenge-shaped biomass distribution - narrow at the
top and bottom and wide in the middle.
A pyramid of numbers represents the number
of organisms at each trophic level - the number of
plants, the number of herbivores, and the number
of carnivores (Figure 6.11). This information can be
troublesome when comparing two different ecosys-
tems - it is not very informative to equate ‘a diatom
with a tree, or an elephant with a vole’ (Phillipson
 COMMUNITIES 153

TOP Parasites Owls Weasels Hyperparasites Shrews Moles

carnivores

Carnivores

Herbivores

Plants

(a) Primary producers (b) Primary producers (c) Plant-parasite food

chain
small individuals
- -- large individuals

--
A-
Tertiary
consumers

-- Secondaryconsumers
Primary consumers
Prlmary producers -
+
t- Hyperparasites
Plantparasites
Primary producers

Figure 6, I I Ecolog~calpyramids. (a) Primary producers are small organlsms. (b) Primary producers are large organlsms.
(c) A plant-parasite-hyperparasite food chain.
Sotow: After Phillipson (1966)

1966: 13). The typical pyram~dof numbers applies depend. Several keystone species have been identified
whenproducersaresmall, as theyare In aquatlc in the wild, butit is not easy to predict whichspecies
ecosystems. In forests, the producers - mainly trees - will be keystone because the connectlonsbetween
arelarge,and a pyramid of thevariousconsumer spec~esIn food webs are often complex and obscure.
levelsperchesona thln base. Inplant-paras~te- For Instance,largecatsactaskeystonepredators
hyperparasite food chains, the pyramid of numbers I S in Neotropical forests. Theylimitthenumber of
Inverted. medium-slzedterrestrlalmammals,whlch In turn
control forest
regeneration. O n BarroColorado
Island, Panama, jaguars (Fdir onla), pumas (Fe/j.r IQN-
Keystone species
cdor), and ocelots (Feii.r pardaiis) have been removed.
Keystonespecies arespecies central to an eco- The populations of big-cat prey - Including the red
system, species upon whlch nearly all other specles coat1 (Na.wu m u u ) , the agouti (Dmypm.tLz twrreptu),
154 COMMUNITIES
andthe paca (Ago//// /1ul27) - are abouttentlmes
hlgher than on CochaCashu,Peru,whereblg cats
stilllive(Terborgh 1988). However,this Increase
may result from natural population varlability rather
than the lack of ~ a g ~ ~and
a r spumas (Wright et 'rL.
1994). Theextreme removal of herblvores and
frugivorous mammals would drastlcally affect forest
regeneratlon,alterlngtree specles composltlon, but
the effectsof modestchanges In densmes are less
clear.
Several examples of keystone specles will be con-
sidered, and then some general Ideas about keystone-
specles removal will be examined.
Keystone predators
The sea otter (Enhydro /u/ri.r) is a keystone predator
pur e x [ d / e t m (Dugglns 1980). A populatton of around
200,000oncethrivedonthekelpbedslylng close
to shorefromnorthernJapan,throughAlaska, to
southernCalifornlaandMexlco(Figure6.12). In
1741,VitusBerlng,theDanlshexplorer,reported
seeinggreatnumbers ofsea ottersonhis voyage
among the Islands of the Bering Sea and the North
Paclfic Ocean. Some f k were taken back to Russia
and soon this new commodity was hlghlyprized
for coats. Huntlng began.In1857,Russia sold
Alaska to theUnitedStates for $7,200,000.Thls
cost was recouped in forty years by selling sea otter
pelts.In1885alone,118,000 sea otterpelts were
sold. By 1910, the sea otter was close to extlnctlon,
with a world-wlde populatmn of fewer than 2,000. It
was hardly ever seen along the Californian coast from
191 1 until 1938.
The inshore marine ecosystem changed where the
sea otter disappeared. Sea tlrchlns, whlch were eaten
by theotters.underwenta population exploslon.
They consumed large portions of the kelp and other
seaweeds. Whiletheotterswerepresent,thekelp
formed a luxuriant underwater forest, reachlng from
the sea bed, where I t was anchored, to thesea surface.
With no otters to keep sea-llrchIns In check, the kelp
vanlshed.Stretches of theshallow ocean floor were
turned into sea-urchrn barrens, which were a sort of
submarlne desert.
Happily, a few pairs of sea otters had managed to
survive In theouterAleutlanIslands ancl at a few
localities along the southern Californian c.oList (Figure
6.12). Some of these were taken to Intermediate sites
in theIJnltedStatesandCanadawherethey were
protected by strict measures. With a little help, the
sea ottersstaged a comeback and the sea urchins
declined. The lush kelp forest grew back ancl many
lesser algae movecl In, along with crustxxuls, squlcis,
fishes, andother organisms. Greywhales migrated
c-loser to shore to park their young I n breaks along
the kelp edge while feeding on the dense concentra-
tlons of anlmal plankton.
Keystone predators sometimes are more effective
wlthlncertalnparts of thelrrange.The sea star
(Pi.iu.Itrr o d w w / . r ) I S a keystonepredator of rocky
Intertidalcommunttles In western North America
(Paine1974). Thls starfish preys primarily on two
mussels - hfyti/m t U / i f ~ T N l r l l l / / . iand h l ~ , / i / /tro.w~/t/r.
/.~
A study along the central Oregon coast showed that
three distlnct predatmn regimes exlst (Menge et crl.
1994).Strongkeystone predation occurs along
wave-exposed headlands. Less strong predation by sea
stars,whelks,and possibly other preclators occurs
In a wave-protected cove.Weak prech tlonoccurs at a '
wave-protected site regularly buried by sand.
Keystone herbivores and omnivores
Somekeystoneherblvores
and
omnlvores are so
domlnant that they help to structure the ecosystems
inwhlchthey live. Beavers, elephants,andhumans
are cases In point.The benver (CL~jtorc.crl/rrr/cw.rc.r)
buildsdams, s o creatlngpondsandraisingwater
tables(Nalman r t d . 1988). The wetter conditions
produce wet meadows, convert streamside forest into
shrubby copp~ceareas. and open g q x In woods some
distance away by toppling trees. The elongated
arra fashioned by a beaver family may exceed 1 km
In length. Hcavers also cause changestothe b~o-
geochcmlcal cha~acter~st~c-s of boreal forest dralnage
networks (Nalman et ' I / . 1994).
Elephants, rhlnoceroses, and other b ~ gherbivores
play a keystone role 111 the savannahs and dry wood-
lands ofAfrlca
(Laws 1970; Owen-Smtth 1989).

Fifure 6.12 Sea otter (Enhydra IUWIJ)
distribution. Circles are its mid-eighteenth-century distribution. Black dots are I
distribution around 1910.-
Sourre: Partly after Ziswiler (1967)
African elephants (Loxodonta afiicana) are relatively
unspecialized herbivores. They have a diet of browse
with a grass supplement. In feeding, they push over
shrubs and small trees, thus helping to convert wood-
land habitats into grassland. They sometimes destroy
largematuretrees by eatingtheirbark. As more
grasses invade the woodland, so the frequency of fires
increases, pushing the conversion to grassland even
further.Grazmgpressurefromwhiterhinoceroses
(Ceratotberium simum), hippopotamuses (Hippopotamus
amphibrus), and eland (Taurotragus oryx) then trans-
forms medium-tall grassland Into a mosaic of short
and tall grass patches. The change from woodland to
grasslandisdetrimentaltotheelephants,which
begintostarve as woodyspeclesdisappear, but
beneficlal to the many ungulates that are grassland
COMMUNITIES 155
~ S
grazers. Over millionsof years, browsing and grazing
by themegaherbivores of sub-SaharanAfricahas
created a mosaic of habitats and maintained a rich
diversity of wildlife.
Some early humangroupswerekeystoneomni-
vores. Their invasion of North America may explain
the mass extinction of megaherbivores (Owen-Smith
1987, 1988, 1989).Hunters may have liquidated the
largestmammalianherbivores.Theseglantspecies
had browsed and grazed, and maintained the grass-
land habitat. The reduction of the megaherbivores,
possiblyinconjunctlonwlthclimatlcchange,led
toforestexpanslonandopen-habitatcontractlon.
Thesehabitatchanges caused manymammaland
bird populations to become reduced and fragmented.
Coupled with additional hunting, loss of prey and
156 COMMUNITIES

carrion for predators and scavengers, andother For simple food chams, the situation is reversed
changes, habitat fragmentation led to the extinctlon (Figure 6 . 1 4 ~ ) Highly
. specialized herbivores or car-
of other species. The human colonizatlon of North nlvores are extremely vulnerable to the loss of their
American may have triggered a cascade of events that sole food source. The koala (Phasco~arrtoscinereus) is an
ultimately caused theextinctlon of the megafauna arboreal folivore. It feeds almost exclusivelyon the
and other species in Plelstocene times. foliage of gum trees (Etlcalyptrrs). Although still wide-
spread,the koala populatlon is controlled In some
areas where overpopulatwnwouldotherwise lead
Removing keystone species
todefoliatlon,thedeath of scarce food trees, and
What happens if a trophic level should be removed anendangerment of the koala population (Strahan
froman ecosystem? Thls I S an enormously difficult 1995: 198). The sabre-toothed cats were one of the
question, to which there are at least four contradic- main branches of the cat family (Felidae) throughout
tory answers (Pimm 1991) (Box 6.1). much of theTertiary.They had enormousupper
The effect of removing specles depends upon the canine teeth and probably specialized in preying on
complexlty of the food web (Figure 6.14). In a com- large, slow-moving, thick-hided herbivores,
such
plex food web, removal of a plant at the bottom has as mastodons and giant sloths. They became extinct
little effect through the rest of the ecosystem (Figure during the Pleistocene, most likely because their prey
6.14a). This is borne out by the limited Impact of was at first thinned by extinctions and finally
American chestnut (Castanea dentata) removal from vanished.
the eastern forests owing to chestnut blight (p. 80). The examples suggest that introduclng generalists
Seven specles of butterfly that feed exclusively on the should have a greater impact on an ecosystem than
chestnut are probablyextinctbutforty-nineother introducing specialists. Correspondingly,introduc-
species of butterflythat also fed on thechestnuts tions into ecosystems containmg generalist predators
found alternative food sources, as didthe insect (which can limitthenumber of intruders)should
predators that fed on all fifty-six butterflies. On the have less of an impact than introductions of speclalist
otherhand, removal of keystone predators o r her- predators.
bivores is not so innocuous an event - a major shock
cascades all the way down the food web and shakes
Contaminating food webs
to lowermost level (Figure6.14b).Kangaroo rats
(D~podomys spp.) are a keystoneherbivores in the Humans are part of food webs. Thelr skills at hunt-
desert-grassland ecotonein North America. They ing, and later farming, enabled them to explolt plants
have a major impact on seed predation and soil dis- andanlmals in amannerquiteunlike any other
turbance. Twelve years after thelr removal from plots organism. They are super-omnivores. Hunting skills,
of Chihuahuan Desert scrub,tallperennialand 10,000 years ago, were equal to the task of drivmg
annual grasses Increased threefold and rodent species largeherbivores to extlnction.Thisdubiousability
typlcal of arid grasslandhad colonized (Brown and has lasted and evolved Into a new, and perhaps more
Heske 1990). Similarly, the cassowary is thought to subtle, form - 6arming has transformed theglobal
be the soledisperser for over a hundred specles of land cover. In large tracts of continents,naturally
woody tropical rain-forest plants In Queensland, diverse plant communities have been replaced by vast
Australia (Crome and Moore 1990). I t usually inhab- expanses of monoculture crops - wheat, maize, and
its large forests. Logglng and habitat fragmentation rice. Ecological efficiency runs at about1 0 per cent, so
have removed the birdfrom several areas, inwhich i t makes more sense for humans to eat plants than to
only small remnants of forest remain. A progressive let herbivores eat plants and then eat the herbivores.
and massive loss of trees I S likely to follow, unless the It makes little sense to let carnivores eat the herbi-
cassowary adapts o r adjusts its behaviour. vores and then eat thecarnivores. Most cultures doeat
 COMMUNITIES 157

l o x 6.1
REMOVING TROPHIC LEVELS - scarcity of herbivores in turn limits the number of
carnivores. If either carnivores or herbivores should
rHREE IDEAS
be removed, the effect upon the community as a
What happens if a trophic level is removed from wholewould be limited becausethey arealready
I community? To answer this question it is helpful kept in check by the plants at the base of the food
:o ask what holds communities together. There are chain. In this model, competition is between herbi-
:hree competing ideas (Pimm 1991) (Figure 6.13). vores andbetweenpredators,and if a species of
either is removed, i t should have consequences for
1
the other species at the same level of the food chain.
The world is green’ hypothesis
Carnivores, which have no predators above them to The world is white, yellow, and
ceep them in check, should keep herbivore popula- green’ hypothesis
:ions low. In turn, the herbivores should have little
impact on plants, which thus thrive. Consequently, This considers the likely effects of ecosystem prod-
removingherbivoresshouldhaveaminoreffect uctivity. ’White’ ecosystems with low productivity,
upon anecosystem,whereasremovingcarnivores such as tundra, contain scant plants that compete
;hould have major effectbecause itwouldallow amongthemselves for limited resources. They do
herbivores to boom. In this model, competition is not produce enoughfood to support more than few a
intense between plants and between predators, but herbivoresandevenfewercarnivores.Removing
not betweenherbivores.Removal of apredator herbivoresandcarnivores,whichcompeteonly
willthereforehavemajorconsequencesforother weaklywiththeirpeers,from‘white’ecosystems
predators. will have little impact. ‘Yellow’ ecosystems, suchas
temperateforestsandgrasslands, have medium
productivity. Enough plant material is produced to
The world is prickly and tastes bad’ support a modest number of herbivores, certainly
hypothesis enough to keep plant numbers in check, but not
Most plants possess defence systems, such as toxins support a largenumber of carnivores.In‘yellow’
and sharp spines, that limit herbivore numbers. The ecosystems,carnivoreremovalswill
have
little

(a) The world is green‘ (b) ‘The world is pricklyand (c) The world is white, yellow,andgreen’hypothesis
hypothesis
bad’
hypothesis
tastes
White’ ’yellow’ ‘CEWl’
1 58 COMMUNITIES

impact, but herbivore removals will have a major of ecosystems. It explain!

nations for certain features
impact
onplants.'Green'
ecosystems,
such as whycarnivoresaresometimesimportantin eco.
Y . 1 I --- "" ""

enough carnivores to keep herbivores in check. In explains why controlling carnivore numbers some
'green' ecosystems, removing carnivores will affect times increases the populationof a prey species,anc
herbivorenumbers,
whichwill
in turn affect sometimesitdoesnot do so. 'White'ecosystems
plants. such as the Arctic tundra, do seem resilient to majol
perturbations. 'Green' ecosystems, such as Yellow.
These
hypothesesare
simplistic, butthe field stone National Park, do suffer from the removal o
evidence tends to favour the 'world is white, yellow, carnivores - herbivorenumbersincreasegreatlj
and green' hypothesis, which offers plausible expla- with an accompanying impact on vegetation.

Carnivores n 0
0 t
r\e 0 0r"\0 080 0
Herbivores

Plants
t
Figure 6.14 Simple and complex food webs - repercussions of removing trophic levels. (a) Removing a smgle plant from
a food web has little impact on a predator, which draws upon a range of food sources. (b) Removing a predator from a
food web creates a 'shock wave' chat cascades down the trophic levels. (c) In a simple food chain, removlng a single plant
species may rnaterlally affect the predator.
Sourre: After Budiansky (1995)

somemeatandherbivoresarethemainsource of or food-chainconcentration.Its inimicaleffects

animal protem. Nonetheless, carnivores do appear in were brought to public notice by Rachel Carson in
some diets. herbook Silent Spring (1962).This book drew
Thehumanexploltatlon of food c h a m IS an attention to the alarming build up of long-lasting
enormous topic. Three issues will be considered here pesticides,mainly DDT,intheenvironmentand
because they impinge on biogeography - biological the damage that they were causing to wildlife and
magnification, the long-range transport of pesticides, humans near the top of food c h a m .
and the long-range transportof radioactive isotopes. DDT is achlorinatedhydrocarbon. It wasfirst
prepared by Othman Zeidler In 1874. Paul Muller
discovered its msecticidal properties in 1939 and,for
Biological magnification
doing so, received the Nobel Prize forPhysiology.
Somesubstances,
Including
pestmdes,may be It was thought to be a panacea, a complete solution
applied In concentrations that are harmless to all but to pest control. By the early 1960s, its perslstence
thepeststheyaredelgnedtoeradicate.However, intheenvironmentandaccumulationinthe food
concentrationsbuildup (or aremagnified) as one cham were becommg apparent. In 1972, after years
organlsm eats another and the pesticide IS fed along of forceful lobbying and petitioning in the United
a food cham. This process IS called biomagnification States, D D T was banned for all but emergency use
 COMMUNITIES 159

by the Environmental Protectlon Agency. The bio- Long-range transport of pesticides

magnlfication of DDT inthe LongIslandestuary,
New York, clearly showstheincrraslngconcentra- Organochlorines (chlorlnated
hydrocarbons)
tlons in moving up the trophlc levels (Figure 6.15). Include the best known of all the synthetic poisons -
One reason that DDT accumulatesalong endrln, dieldrin, lindane, DDT, and others. They are
food
wldely used as biocides. Polychlorinated biphenyls
chains I S that t t IS firmly held in fatty animal tissues.
Some heavy metals are also stored in body tissues and (PCBs) are used in plastics manufacture and as flame
are subject to biomagnlfication. In the Alto Paraguay retardants and insulating materials. A very worrying
River Basln, Brazil, large quantities of mercury, used development is that h1ghlevelsof organochlorines
in gold mining, arc dispersed directly into the are recorded in Arcticecosystems,wheretheyare
atr and
the rlvers runningintoPantanal, a wildlife reserve biomagnitied and have lnlmtcal effects on consumers.
The organochlorinesarecarrrednorthwardsfrom
(Hylanclrr e r a / . 1994).Local, commercially important
agriculturalandindustrlalreglons
catfish ( P . r e / t ~ / O ~ / ' / t ~ , . ~m f ox ~~~ ~/ ~/ / . rhad
) a mercury In theair.They
content above the limit for human consumptlon, and then enter Arctic ecosystems through plants and start
slgnificantlyabovethenaturalbackgroundlevel. an upward journey through the trophic levels.
Mercury content In bird feathers PCBs and DDT are the most abundant residues rn
also indicated
biomagnificatlon. No statistlcally significant accumu- peregrine Falcons ( ~ & / ~ - ~ ~ e ~ e s t(D.. / l J.
/ / tThomas
~.) et a/.
latlonofmercury 1992). They reach average levels of 9.2 and 10.4
was found In soil andsediment
samples.Evidently,mercuryoriglnatingfromthe &lg, respectively. Theseconcentratlonsaremore
goldmining process is more readilyabsorbed thantentuneshigherthanotherorganochlorines.
by
organismsthanmercurynaturallypresent They are also found in polar
in soil bears (Ur.rl4.r rnurititnw)
mlnerals. (Polischuk et a/. 1995). A wide range of organochlo-
Ditches along busy roads are liable to pollution by rine pesticides andPCBsweremeasured In muscle
heavy metals. In Louisiana, UnitedStates,anrmak tissue and livers of lake trout (Salvelinlrs namaymsh)
and plants living in ditches have accumulatedcad- and Arctlc grayling(Thyrnalhrsarrtrrus) from Schrader
mium and lead (Naqvi et &/. 1993). In the red swamp Lake, Alaska (R. Wilson et a/. 1995). PCBs are
crayfish (Procan/huvts d r r k i i ) , thecadmium recorded in martneArcticecosystems,too.Samples
level
was 32 tlmes that In the water, and the lead level 1 2 were collected near Cambridge Bay, at Hall Beach,
tlmesthat In thewater,glvingbioaccumulation and at Wellington Bay, all in Northwest Terrltorles,
factors of 5.1 and 1.7, respectlvely. Canada(Brtght et n / . 1995). Organismsstudied
Included clams (Mya trlmzta), mussels (Mytilzts
e d / r / i j ) , sea urchins (Stron~y/orentrot/{~. droeha'hiensis),
Long-range transport of radioactive isotopes and four-horned sculplns (hIyoxocz/hal/rs quadrzrornrs),
Although southern and temperate blologlcal systems whlchare fish. Some of the high concentrations in
have largely cleansed themselves of
radioactive thesesampleswereattributable to local organo-
fall-out
depostted
durlngthe 1950s and 1960s, chlorine sources, but long-distance sources were also
Arcticenvironments have not (D. J. Thomas et al. implicated.
1992). Lichens accumulateradioactivtty more than
many other plants because of thelr large surface area
andlonglife-span; the presenceandperslstenceof F R O MB A C T E R I AT OB L U EW H A L E S :
radioisotopes in the Arctic are of concern because B I O D I V E R S I T Y
ofthe lichen-reindeer (Ran<qj/ir /rrr~/~/r~/ts)-human
ecosystem. Biological diversity, or biodiversity for short,
incorporates three types of diversity - genetic, habi-
tat, and species. Genetic diversity I S the variety of
E
3
C
W
T
.-m
C
r
U
._
C
 COMMUNITIES 161

information (genetic characteristics) stored in a gene Table 6.3 The diversity of living things
pool. Habitat diversity IS the number of different
habitatsin a given area. Species Number
diversity is the Group of
number of species in a given area. It is also called species
species richness and species number.
Guestimates of the total number of species cur- Insects 75 1,000
rentlylivingontheEarth vary enormously.They Other animals 28 1,000
range from 4.4million to 80 million. The number of
Higher plants 248,400
Fungi 69,000
known species is about 1,4 13,000. Therefore, the low
Protozoa 30,800
estlmate of 4.4 million would mean that 32 per cent Algae
of all species are known to date; the figure drops to a Bacteria and similar forms 4,800
mere 2 per cent for the high estimate of 80 million. Viruses 1,000
Thetotal species diversitydisguises enormous
differences between group of organisms (Table 6.3). Source: From data in E. 0. Wilson ( 1 992)
Insects are by far the most numerous group on the
planet.Totaldiverslty also disguises threeimpor- Species-area curves
tantgeographical diverslty patterns - species-area
The Increasing number of species with increasing
relationships, altitudinal andlatitudinal diversity
area I S described by a species-areacurve.Figure
gradients, and diversity hot-spots.
6.16 is a species-area curve for plants in Hertford-
shire, Enghnd. To construct the curve, plant species
Species-area relationships records from10-kmgrid squares were grouped
Into successively largercontiguous areas, untilthe
Count the species in increaslngly large areas, and the entire county was covered. Figure6.16a shows the
species diversity will increase. This, the species-area data plotted on arithmetic co-ordinates. The result
effect, is a fundamental biogeographical pattern. It is a curvilinear relationship. The same data plotted
applies to mainland species and to island species. on logarithmlc co-ordinates (Figure 6.16b) produces

y = 480.581 + 0.41 7x log y = 2.21 3 + 0.265 log x

r 2 = 0.589 r2 = 0.836

I I I I I I I 2.0
0 1,500
500 1,000 0 1.o 2.0 3.0
Area, x (krn2) Log area, x (krn2)
162 COMMUNITIES

a linear relationship between specles recorded and The z-value (0.27) indicates that, for every 1 kmz
area. The line is described by the equation increase inarea,an extra 0.27 plant specles will be
log S = log c + z log A
found.
Species also increase with area withm Island groups "

where S is the number of species, A is area, c is the - large islands house more species than small islands.
intercept value (the number of species recorded when For amphibians and reptiles (the herpetofauna) living
the area is zero), and z is the slope of the line. This on theWestIndian islands, therelationship, as
equatlon may also be wrltten as depicted on Figure 6.17a, is
S = '.A' S = 2 . 1 6 ~ ~ ' ~ ~
For the Hertfordshire plants, the equation is
The line described by thls equation fits the data for
S = 2.21A'''' Sombrero,
Redonda, Saba, Montserrat,
Puerto Rlco,

/
Trinidad

Puerto RicG
/
__
Hispaniola
'Cuba
7)
6 1.5 4 -w
Jamaica

log S = 0.366+0.37 log A

(S = 2.16A037)
f
M
r' = 0.92
Sombrero
2
0 !: I I 1 I I
2 0 1 3 4 5
Log area, A (km2)
". 3.0 - (b)
m
.-aU,
%
2.5 -
-m0.
L
0
L

2 2.0 - log S = 2.1log

80+0.175 A
5 Stronsay 0 (S = 151S A 0 ' 7 5 )
M r' = 0.39
2 1.5 I I 1 I I
-1 0 3 1 2 4
Log area, A (krn')
 COMMUNITIES 163

Jamaica,Hispanlola,andCuba very well.Trlnidad Important for plants.Fourteenhabltattypeswere

lies well abovetheline. It probablycarriesmore Identified (Table 6.4). The results show strong pos-
species than would be expected because it was joined Itlvecorrelatlonsbetween spec~esper Island,Island
to South America 10,000 years ago. For plant species area, andhabltatdiverslty(Figure 6.18). Habltat
onaselectionofScottlsh Islands, therelatlonship diverslty Itself correlatedwith Isl;md S I X . Atech-
(Figure 6.17b) is nlque called path analysls was ~lsedtodistlnguish
the effectofIsland sizeonspeciesdiversityfrom
S = 151.51A"1X
the direct effect of habltat diversity. The sum of the
Alargenumberofstudies have establishedthe direct effects(area actlngthrough areaalone)and
validity of this klnd o f relat~onsh~p. Indirecteffects(areaactlngthroughhabltatdiver-
As a rule, the number of specles living on Islands sity) of area were almosttwlcethe overall effect
doubles when habitat areaIncreases by a factor of ten. of habltat diverslty on spec~esdiversity. Thls finding
For islands, values of the exponent z normally range strongly
suggests a directrelat~onshipbetween
from 0.24 to 0.34. In malnland areas,z-values nor- Island area and speciesdiverslty, ~ndependent of
mally fall wlthln the range 0.12 to 0.17 (the value habitat diverslty.
of 0.27 for Hertfordshlreplants mayresultfrom
thehighgeodiversity of thatcounty).Thls means
Diversity gradients and hot-spots
thatsmall areas containalmost as many speciesas
largeareas, butsmall Islands contaln fewer species
The latitudinal diversity gradient
thanlarge Islands. The differencemaybepartly
attributableto relatlveIsolation of Islands, which Many more spec~eslive In the tropc-s than live I n the
makes colonization more difficult than on mainlands. temperate reglons, and morespecies live In temperate
reglons than live in polar reglons. In consequence, a
latitudinal species diversity gradientslopes steeply
The habitat diversity and area-alone
away fromatroplcal'highdiversityplateau'. The
hypotheses
cliverslty gradient I S seen In virtually a l l groups of
A crucialquestion is why there is such agood organism. An example I S the species richness gradient
relationship between area and species diverslty. The of American mammals (Figure 6.19). The diverslty
answer is notslmple.Two rival hypotheses have falls fromatropicalhlgh of about 450 mammal
emerged (Gorman 1979: 24). The habitat diversity specles to a polar low of about 5 0 mammal species.
hypothesissuggeststhat largerareas have more Why are there so many spec~es (and genera, and
habltats and therefore more species. The area-alone families,andorders)inthetropics? Or, conversely.
hypothesis proposes thatlarger areas should carry why are there so few species in temperate and polar
more species, regardless of habitat diverslty. A study regions?Thesearefundamentalquestionsinhio-
of dicotyledonousplant speciesonforty-five un- geography and ecology and have exerclsed the mlnds
inhabited,unimproved, small Islands off Shetland of researchers for over a century. There is no shortage
Mainland,Scotland, plus twosimilar
headlands of suggested answers (Table 6.5). A survey of twenty
treated as islands, sought to test the two hypotheses hypotheses suggests thateach one contains an element
(Kohn and Walsh 1994). Species lists were complied ofclrcularlty or else IS not supported by sufficient evi-
during a systematictransect search coverlng each dence (Rohde 1992). Hypotheses flawedby clrc-ular
Island. A second species list was derived by placing reasonlngincludeblotlchabltatdiverslty,competl-
50-cm X 50-cm square
quadrats
randomly on tion, niche width, and predation. Hypotheses that 1ac.k
twenty-two Islandssufficlentlyvegetated for a rea- an empirical base include: climatic stability, climatic
sonable number of samples. Habltats were classified varlability,
latitudinal
ranges,
arm,
geodiversity,
accordingto physical characteristlcsassumed to be and prmary production. It I S possible that none ofthe
 164 COMMUNITIES

Table 6.4 Habitat classificationused in Shetland study

Class Type Number

~ ~~~ ~ ~ ~ ~~~~ -

Rocks Sea cliff 1

Scree 2
Boulder field 3
Pasture General, dry 4
Rocky pasture 5
Wet pasture: Damp depressions, runnels 6
Moss-dominated wet ground 7
Mud 8
Bog 9
Open standing water 10
Stream 11
Grass cliff 12
ShinLgL 13
ShinLgle
Sa n Son 14

Total number of habitats 14

-
Source: After Kohn and Walsh (1994)

w
* ** I'i
60 (b'
0

* *

* *
30
IC) * :
** y o -

l : 1
0.1 0.01 10 100 0
Island area (hectares)
 COMMUNITIES 165

500 Perissodadyla
..........
........... Primates
Xenarthra
400 Didelphimorphia
0 Chiroptera
rn
Carnivora
2
c
300 Artiodactyla
.-
U
L

.-$
200
VI

100
. .. .. .. .. . .. . . . . .
......
..............
0
70 60 50 40 30 20 10 0 10 20 30 40 50
N (degrees) Latitude 5
Figure 6.19 Latitudinal diversity gradient mammal species richness in the Amencas. The species are grouped into Orders.
Two Ordersareomrttedbecausethey contaln too few species. The Microbiotheria contains one specres that ranges
from 35"s to 45"s. The Paucituberculata contain threespecres found In bandsfrom 15"N to 20"s andfrom 30"s to
45"s. Notice that species richness is fairly constant on the 'tropical species rrchness plateau' and declines steeply with
increasmg latltude outslde the troprcs.
Source: After Kaufman (1995)

expenditure.Abioticfactorsaremostlikelyto be species distributions, and allow many specles to live

severely limiting in polar regions and relax towards in the tropics. Ultimately, the interplay of abiotic
the equator. Contranwlse, biotic interactions areless and biotic factors generates the latitudinal gradient
limiting In polarregionsbecause species diversity in species diversity. Latitudinal diversity gradients
is low. They become severer towards the equator. As occur in genera, families, and orders, as well as in
I t becomeseasier tocopewithabloticconditlons, species (Box 6.2).
speciescan devotemore resources toInteracting
with other species, that is to competing. By these
Diversity hot-spots
arguments, In passing along the gradient from the
abioticallystressful poles tothemoreablotically Superimposed on latitudinal and altltudinal trends,
congenial tropics, biotic lnteractions should become are diversity hot-spots. These are areas where large
increasingly important in limiting species distribu- numbers of endemic species occur.Eighteenhot-
tions and influencingspecies diversity. spots have been identified globally. Fourteen occur in
Ageneralexplanation for latitudinalgradients tropicalforestsandfourinMediterraneanbiomes.
emerges from these ideas (Kaufman 1995). Abiotic These hot-spots contain 20 per cent of the world's
conditions limit diversity by setting the higher lati- plant species on 0.5 per cent of the land area. All are
tude boundaries of species distributions, and permit under Intense developmentpressure.
only a few specles to live near the poles. Biotic inter-
actions become limiting where abiotlc conditions are
more favourable, set the lower latitude boundartes of
166 COMMUNITIES

Table 6.5 Some factors thought to influence species diversity gradients

factor Rationole
.___ .~

History More time allows more colonization and the evolution of new
species. In polar and temperate regions, diversity was greatly
reduced during the ice ages and is now building up again
Climate The warm, wet, and equable tropical conditions encourage a
smaller niche breath, and therefore more species, than the colder
and highly seasonal conditions elsewhere
Climatic stability Tropical climatic stability is conducive to species specialization
(smaller niche width) and therefore more species
Habitat heterogeneity The greater the habitat diversity, the greater the species diversity.
Forests contain more niches than grasslands. Tropical forests contain
more niches than any other biome
Primary production In food-deficient habitats, animals cannot be too choosy about their
prey; in food-rich habitats, they can be selective. So food-rich
environments allow greater dietary specialization and smaller niche
width
Primary productionstabilityHabitats with stable and predictable primary production should
allow greater dietary specialization (smaller niche width) than
habitats with more variable and erratic primary production
Competition Competition, which is most intense in the tropics, favours reduced
niche width
Predation Predation reduces competitive exclusion. Predators are therefore
‘rarefying agents‘, reducing the level of competition between their
prey species
Disturbance Moderate disturbance mitigates against competitive exclusion
Energy Species richness is limited by the partitioning of energy amon
species. In the energ -rich tropics, there is more energy to ‘di$ out’
and it can be spreaJaround a larger number of species than in
temperate and polar regions
Latitudinal range size Range size tends to increase towards the poles (Rapoport’s rule), so
fewer species can be packed into a given area
Area The tropics cover more area than any other zone, which stimulates
speciation and inhibits extinction (Rosenzweig 1992, 1995)

Diversity
change extinction
the rate exceeds the speciation
rate. A
point often overlooked in discussions of mass extinc-
Change in species diversity depends upon gains and tions is that they could result from a reduction in the
losses from theglobaldiversity pool. Species arespeciation rate whiletheextinctionrate stays the
added by speclatlon and lost by extinction. Plainly, same - they may not slmply result from an elevated
global biodiversity will rise whenever the speciation extinction rate.
rate exceeds the extinction rate; and it will fall when Complications arise whenthebiodiversity of a
 COMMUNITIES 167

Polar Pollution,
overhuntlng, overfishing, the wildlife
zone trade, and above all, habitat destruction (land cover
Ablotic change - the conversion of wild habitats for farmlng,
Biotic
/ factors fuel extraction, industry, and residential land) are the
Temperate main problems. The main habitats lostare tropical
Ablotic zone
rain forests, wetlands, and coral reefs.
Biotic
factors
Does biodiversity matter?

Although it is rather envlronmentally hostile to do

Abiotic so, I t is worth asking if loss of biodiversity matters.
-, --
factors --
Tropical Is the b-word slmply a buzzword, or is there some-
zone
Biotic
factors
F i g w e 6.20 Controls
on
blotic diversity In polar,
temperate, and tropical zones.
Suurce: After Kaufrnan ( 1 995)
region is considered. In thls case, lmmlgration and
emigratlon of species enter the equation. The bio-
diversity of the Brltish Isles will depend upon the
balance between the speclation rate and lmmlgration
rate on the one hand, and the extinctlon rate and the
emigration rate on the other hand. These relation-
ships are analogous to changein populations (Chapter
4). Speclatlon IS equivalenttobirths,extlnctionto
deaths,andlmmlgratlonandemlgratlonapplyto
species rather than to individuals.
Why all the fuss?
It is the fashion to bandy figures aboutextinction
rates in the present wildlife holocaust. Extlnction is
said to be running at 25,000 tlmes the natural rate.
By the year 2000, onemillionanimalsandplants
will have been driven to extinction, and the extlnc-
tlon rate will have soared to several species per hour!
By 2050, half the species alivetodaycould have
gone. And, as they say, extinction is forever.
Although some ecologists are stepping back from
these hyperbolic estlmates, there is little doubt that
extinctlon I S runnlng very fast at present and will
continuetodo so. The chief culprit is humans.
thingimportanthldingbehindthe biodiversity
rhetoric? This questlon may be answered by asking
twoquestions.First, IS the
currentbiotlc crlsis
unique? Second, what stands to be lost?
The only yardstlck for comparingthecurrent
extlnction rate I S the rate of species loss during the
mass extinctionsthat have occurred through geo-
logical time. Several past mass extinctions,though
described as ‘catastrophic’, took hundreds of thou-
sands of years. The Plelstocene mass extinctions were
somewhat speedier. Some evidence suggests that the
current loss of biodiversity I S ‘thebiggest mass-
extinction of them all’ (Leggett1989).
A loss of biodiversity should perhaps be deplored
on moral grounds -most people would accept that all
creatures have a right to exist. In addition, and this
seems to carry far more weight with decision-makers,
a loss of diversity is accompanied by a loss of genetlc
resources (genes, genomes, and genepools), medicinal
substances, and potential foodstuffs. It is surprlsing
how dependent the
pharmaceutical industry is
upon the natural world. In the United States, of all
prescriptlons dispensed by pharmacles, 25 per cent
arederivedfrom plants, 11, per centfrommicro-
organisms, and 3 per cent from anlmals. As for food-
stuffs, the world currentlydepends onjust twenty
species to provide 90 per cent of its food. Three
species - wheat, maize, and rice - account for over
half the world’s food productlon. The production of
these foodstuffs is biased towards cooler climates and
the cropsarenormallysown as monoculturesthat
brmg attendant problemsof sensitivlty to disease and
attacks frominsects andnematodeworms. Many
168 COMMUNITIES

Box 6.2
L A T I T U D I NDAI LV E R S I T Y decreasing
in strength families,
by
genera, anc
G R A D I E N T S , IN G E N E R A , species.
Just
a few body plans can
survive
under
tht
FAMILIEA S ,NODR D E R S severe abiotic
stresses of high
latitudes,
and thosc
bodyplanstend tobegeneralized. So, abiotic
Animalswithingenera, h i l i e s , andordersshare a factorsactabsolutely on diversity, limiting thc
L ~ " - "1 1 * ~ 11 . . I 1 t ~.~~ "3 L :"
LSIC D O ~ Yplan (Dauplan) ana many ocner cnarac-
\
IlUIIIUTr 01 U W Y pldllb, alIU, UCLdUbC a S P t X l C S CdllllUt
~~~ ~ ~ "" ""

ristics that constrain their distribution and diver- exist where its body plan cannot exist, the number
ty (Kaufman 1995). Members of the same genus of species. The seventeen bat species found north
arealikeinmanyparticulars of theirmorphology, of 45"N belongtoonefamily body plan - thc
physiology,behaviour,andecology. The grass voles,Vespertilionidae.Likewise,theeightxenarthrar 1

- ..p-.- .
."YL.". ..V"C.. "I ,," -11 "\-."..a C" ...
L "L... y
grasslands.They feed ongrassesandsedges,and Dasypodidae.Conversely,
biotic
factors have :a
have ever-growing cheek teeth with a sharpy angu- bottom-upinfluenceonthetaxonomichierarchy
lartriangularpatternonthegrinding surfaces. and their primary effects are felt at the
species level
Members of the same Order share a basic suite of They do not limit the numberof species per se, bu t
characteristics. All bats(Chiroptera), for instance, they do limit each species individually by influenc
. . ..".. .
lare t r a m associated wlth fllght. Body plansare
-. I .
Ing population aynam~cs ana nlcneI. , wlacn.
. I , ?I.o n:
m c 1

not limited by biotic interactions. Rather, they are factorslead to specializationandspeciespacking.

constrained by abiotic factors- a body plan suitable Bioticfactorsinfluencemacrotaxonomicdistribu-
for an aquatic existence is hardly suited to life in tion only if the body has a feature that affects the
deserts (there are no desert-dwelling seals!). Abiotic interactions among species.
factors have a top-down influence on the taxonomic Each species and body plan trades off a resistance
I-.- -IL
kerarcny. wt-.- . - a .~~.~
1 ney mnuencegeograpnlcru
J.--:L..
. ~ ~ ~ - ~ . I L . - I
alstrmu- LU ~UIULIL
---".."
-L:--:- p c a x u c a -aL
:."
ILJ
,.I.,
plaL-luux ,A,
cugc -..A
alu

nns mnsc ntrnndv at the level of orders. fnllowed to biotic Dressures at its eauatorial-most edge. The

Latitudinal range of body plan

"H
-4
" I
I
'Abiotic
constraints

Tropical
Temperate Polar Tropical Temperate Polar Tropical
Temperate Pola

Tempero-polar Intermediate Tropical

body plan body plan body plan

'igure 6.21 Latitudinal variations in abiotic pressure and biotic pressure acting on species of three hypothetical body
-
Aans a tempero-polar body plan, an intermediate body pian, and a tropical body plan. The horizontal lines in the
;haded areas represent latitudinal species ranges within a body plan.
fourre: After Knufman ( 1995)
 COMMUNITIES 169

:rade-off strategies of species and body plans pro-
h c e different latitudinal
distributions
(Figure
5.21). Abioticlimitationsoperatetoasignificant
#ectonly outsidethetropics.Bioticlimitations
lave a nearly constant effect in the tropics and then
:ail off towards either pole. Combined, abiotic and
i o t i c factorslimit species distributionswithina
mdy plan, and influence the body plan itself. Body
dans occupy larger latitudinal ranges than species,
because most body plans consists of m a n i species.
Body planstend to be limitedonly by abiotic
factors. Normally, they straddle the tropics and are
limited by abiotic factors ateach polar-most edge of
their distribution. On the other hand, species are
potentially limited by abiotic and biotic pressures.
Therefore,theyhavereduceddistributions,com-
pared with their body plans, owing to the trade-offs
in adapting both these pressures.

alternative food sourcesareavailable,thoughthe centimetrestotheentireblosphereandecosphere.

menu will shortenas blodiverslty drops. Organlsms have roles withln a community. Some are
producers, making biomass from energy and mineral
resources. Gross primary production is the amount
What can be done to safeguard
of biomassproducedinaunittime.Netprimary
biodiversity?
production IS gross primaryproduction less the energy
Four things, princlpally -take conservation measures,burned by primary producers. The primary produc-
counter the wildlife trade, study diversity, and sell a tioniseaten by consumers.Macroconsumersare
greener envlronmental ethlc. Conservation IS a vast herbivores,carnivores,and top carnivores.Micro-
topicand beyond the scope of thisbook.Alarge consumers are decomposers and detritivores. Energy
amount of time and effort goes Into saving speciesflows (e.g. through
ecosystems. Biogeochemlcals cycle
elephants and whales), savrng habitats (e.g. wetlands around ecosystems,mineralization of deadorganic
In Irlan Jaya andZambia),savlngforests (e.g. in remainsproviding foodforrenewed plant growth.
Ecuador, Costa Rica, andthe Philippines), and saving Feedingrelationshipswlthincommunltiesproduce
islands (Madagascar and the Galipagos). Three main food chains or food webs. Grazlng food webs involve
conservation measures are currently being taken. First, plant + herbivore + carnivore + top carnivore
the establishingof protected areas and natlonal parks. sequences. Decomposer food webs lnvolve commmu-
At present, 5 per cent of the land area IS occupied by tlon spirals.Food c h a m have characteristic ‘pyramids’
nature reserves, national parks, wildlife sanctuaries, of biomass, energy, and numbers. Keystone species are
and protected landscapes. Second, the setting up of crucial to certain communities. Take them away and
IUCN/WWF (International Union for Conservation thecommunitycollapsesorchangesdrastically.
of NatureandNaturalResourcesWorldWildlife Humans have contammated many food webs, largely
Fund) Centres of Plant Diversity. Some 250 sites and becausetoxlcsubstances becomeincreasmglymore
regions of high plant diversity would protect 90 per concentrated as they move up the trophiclevels. Bio-
cent of world plant species. Third, the designating diversity is apressmgconcern. It has aconsistent
of more UNESCO BiosphereReserves. relationshlp wlth Increasing area, as seen in specles-
area curves. There is a latitudinal diverslty gradient
- diversity decreases towards thepoles from a tropical
SUMMARY high-diversity plateau. There are also several diversity
hot-spots.Biodiversityiscurrentlyfalling,owlng
Communitiesarecollections of interactingpopu- largely to habitat fragmentation. The loss of biodiver-
lations livmg In a particular place. Ecosystems are sity appears to be real, despite all the hype, and mea-
communities and their physical environment. Com- sures are in hand to mlnlmize losses during the next
munities and ecosystems rangesize in from afew cublc century.
170 COMMUNITIES

E S SQ
AUYE S T I O N S Reaka-Kudlka,
M. L., Wilson, D. E.,Wilson,
and E.
0. (1 997)Biodivevsity 11: Understanding and Protectrng
1 What are the
similarities and O w Biological Resources, Washington,
DC:
National
differences between grazing food Academy Press.
chains and detrital food chains?
2 H~~ are biogeochemical Rosenzweig, M. L. (1995) Species Diversrty in Space and
cycles? Unwerslty
Cambridge
Time, Cambrldge: Press.

3 Why does it matter if global
biodiversity falls?
F U R T H E RR E A D I N G
Archibold, 0. W . (1994) Etnlogy of WorldVegetation,
New York: Chapman & Hall.
Chameides’ w’ L‘ and Perdue’ E’ M’ (1997)
BiogeochenrrralCycles: A Computer-Interactwe Study o f
Earth System Srrence andGlobalChange, New York:
Oxford University Press.
Schultz, J. (1995) The Ecozones of the World:The
Ecological Divlszons of Geosphere,
the Hamburg:
Springer.
Szaro, R. C. and Johnston, D. W. (1996) Bzodiverszty
in Managed LandsLzpes: Theovyand Practzce, New York:
Oxford University Press,
Wilson, E. 0. (1992) The D~vevsity of Lye,
Cambridge, Massachusetts: Belknap Press of Harvard
University Press.

Hochberg, M. E., Colbert, J., and Barbault, R. (eds)

(1996) Aspects of the Geneszs andMairrtenan~v of
Biological Diversity, Oxford: Oxford Universlty Press.

Jeffries, M. L. (1997) Biodiversrty andConsewation,

London and New York: Routledge.

Lawton, J. H. and May, R. M. (eds) (1995) Extinction

Rates, Oxford: Oxford University Press.

Morgan, S. (1995) Ecology and Enmronment: The cydes

of Lye, Oxford: Oxford Unwersity Press.

Polis, G. A.andWinemiller,K. 0. (1995) Food

Wehs: Integration o f Patterns and Dynamrcs, New York:
Chapman & Hall.

Quammen, D. (1996) The Song of theDodo: Island

Brogeography in an Age of Extinctions, London:
Hutchinson.
 7

COMMUNITY CHANGE
Communities seldom stay the samefor long - community change i s the rule. This chapter
covers:

communities and ecosystems in balance

communities and ecosystems out of balance
human-inducedcommunitychange
communitiesand global warming

As Nature abhors a vacuum, s o the biosphere abhors environment In sucha way as toenticefurther
bare ground. Land stripped clean by fire, flood, colonlsts, untilastablecommunity evolves whose
plough, or any other agency I S soon colonlzed by life. equilibrium endures. The full sequence of change is
Whathappens next is thesubject of considerable called vegetation succession.
debate. Proponents of two main schools of thought Clements believed in the idea of monoclimax. He
have their ownviews on the matter. Theseschools may argued that a prisere was a developmental sequence
be dubbedthe‘climatic climax and balancedeco- that, for given climatic conditions, would always end
system’ school, and the ‘disequilibrlum communltles’ in a reasonably permanentstage of succession -
school. climaticclimaxvegetation.However, he realized
that
climatic climax formations contain
patent
exceptionstothis rule. He termed these aberrant
T H EB A L A N C E OF N A T U R E : communitles proclimax states. There are four forms
EQUILIBRIUMCOMMUNITIES of proclimax state (Figure 7.1): subclimax, disclimax,
preclimax, and postclimax (anticlimax IS somethlng
Climatic climax very different):

Thls Influential school of thought had Its heyday In 1 Subclimax is the penultimate stage of succession
rhe1920sand1930s.It stressed harmony, balance, In all completeprlmaryand secondary seres. It
order,
stability,
steady-state,
and
predictability persists for a long t m e but IS eventually replaced
within plant and animal communities. Itschief expo- by the climax community.In eastern North
nent was Frederic E. Clements, anAmerican botanist. America, an Early Holocene coniferous forest even-
Clements(1916) reasoned thatthe first plant tually gave way to a deciduous forest. Clementsians
colonists are good at eking out a living under difficult would therefore conclude that
the coniferous
conditions. By altering the local environment, they forest was a subclimax community.
pave the way for further waves of colonlzatlon by 2 Disclimax(plagioclimax) partially or wholly
specles with a less ploneering spirit. And the process replaces or modifies thetrueclimax afteran
continues, each new group of species changmg the environmental disturbance. Most of the so-called
172 COMMUNITY CHANGE

natural grasslandsin the

British Isles lying Clements's monoclimax hypothesiswas superseded
below 500 m are maintained by cattle and sheep by a polyclimaxhypothesis (e.g.Tansley 1939).
grazing. If grazing were to cease, the grass- According tothe polyclimax hypothesis, several
landswouldrevert to a deciduous forest. In the different climax communitles could exist in an area
arid and seml-arid United States, vegetation suc- with the same regional climate owing to differences
cession around some ghost towns IS permanently in soil moisture,nutrient levels, fire frequency,
altered by anannual grass - cheatgrass (BYOW~J and so on. The climax-pattern hypothesis (R. H.
tectorrrm) - Introduced from theMediterranean Whittaker 1953) was a variation on the polyclimax
(Knapp 1992). theme. As In the polyclimaxhypothesis, I t argued
3 Preclimaxandpostclimax are caused by local that natural communities are adapted to all environ-
conditions, often topography, produclng climatic mental factors. But i t suggested that there is a con-
deviations from the regional norm. Cooler or tinuity of climax types that grade Into one another
wetter conditions promote postclimax vegetation; along environmental gradients, rather than forming
drier or hotterconditionspromote preclimax discrete communitles that change throughvery sharp
vegetation. In theGlacier Bay area, south-east ecotones.
Alaska, theclimaxvegetatlon is spruce (Sitka
sztchensisthemlock (Tsrqa spp.) forest, butat
Balanced ecosystems
wetter sites bog mosses (SphagnnnL spp.) dominate
wlth occasional shore pine (Pinus contorts) trees to The idea of balanced ecosystems emerged in the
form a postclimax vegetation (see pp. 174-7). 1940s and persisted until the late 1960s. It did not

- Disclimax or
plagioclimax
community
-

1 ,
re:,
Partial Plagiosere Subsere
displacement
(e.g. grazlngl
Climatic climax 4
7 .community 1-4
I
I

displacement (secondary
Subsere
Subsere
(e.g. felling, displacement)
cultivation)
Arresting
Bare soil or
relic community
Prisere

Bare inorganic

Figure 7. I Types of climax comrnunltles. The subclimax community is also called 'deflected succession'
S w w : After Eyre ( 1 963)

differ radically from the idea of Clements's endurlng
climaxes. But it didswitchtheemphasis from
plantformationsto ecosystems and related Ideas of
energy flow, trophic levels, homeostasis, and r- and
K-selectlon. An ecosystem was defined as abiotic
community together wlth its immediate life-support
system (soil, water, and air) (Tansley 1935). During
the next four
decades, the ecosystem idea was
promotedand refined by Raymond L. Lmdeman,
G. Evelyn Hutchinson, and,
most persuasively,
Eugene P. Odum. To Odum, each ecosystem has a
strategy of development that leads to mutualism and
co-operationbetweenIndividual species; in other
words, that leads to a balanced ecosystem. The Gaia
hypothesis is, in part, an extenslon of this Idea. I t
sees an overall homeostasis in the ecosphere, sug-
gestlng that life and its supportlng envlronment are
a globally balanced ecosystem (e.g. Lovelock 1979).
From bare soil to climax
Threemechanisms are thoughttodrive succession
- facilitation, tolerance, and inhibltlon (Horn 1981).
Researchers are divlded over the relatlve importance
of these mechanisms, so there are three main models
of succession.
Facilitation model
The pioneer species make the habitat less suitable for
themselves and more suitable for a new round of
colonists. The process continues, each group of species
facilitatlng the colonization of the next group. This
is the classical model of succession, as expounded
by HenryC. Cowles andClements. According to
Clements, vegetatlon successton involves a predeter-
mined sequence of developmental stages, or sere,
thatultimately leads toaself-perpetuating,stable
community called climatic climax vegetation. H e
recognized six stages in any successional sequence:
1 Nudation - an area is left bareafter a major
disturbance.
2 Migration - species arrive as seeds, spores, and so
on.
COMMUNITY CHANGE 173
3 Ecesis - the plant seeds establish themselves.
4 Competition - the established plantscomplete
with one another for resources.
5Reaction - the established plantsalterthelr
environment and so enable other new species to
arrlve and establish themselves.
6 Stabilization - after several waves of coloniza-
tlon, an enduring equilibrium is achleved.
Reaction helps to drive successlonal changes. O n a
sandy British beach, the first plantto colonize is
marram-grass (Ammt,phila arenarza). After a rhlzome
fragment takes hold, I t produces aerial shoots. The
shoots Impede wlnd flow and sand tends to accumu-
late around them. The plant is gradually burled by
the sand and,to avoid 'suffocation', growslonger
shoots.Theshootskeepgrowingandthemound
of sand keeps growmg.Eventually,a sand dune is
produced.This is then colonized by other species,
including sand fescue (Festuca ruhra var. arenaria),
sand sedge (Carexarenarza), sea convolvulus (Calys-
tega soldanella), andtwo sea spurges (Ellphorbia
paralias and EzIfihorhza portlandicz), thathelpto
stabilize the sand surface.
Tolerance model
According to this model, late successional plants, as
well as early successional plants, may invadein the
lnltial stages of colonization. In northern temperate
forests, for example, late successional species appear
almost as soon as early successlonal specles in vacant
fields. The early successional plants grow faster and
soon become dominant. But late successional main-
tain a foothold and come to dominate later, crowding
outthe early successional species. Inthismodel,
succession I S athinnmgout of species originally
present, rather than aninvaslon by laterspecieson
ground prepared by specific pioneers.Any species
may colonize at the outset, but some species are able
tooutcompeteothersand come todominatethe
mature community.
174 C O M M U NCI THYA N G E
Inhibition model
Thismodel takes account of chronicandpatchy
disturbance, a process that occurs when, for example,
strong winds topple trees and create forest gaps. Any
species may invade the gap opened up by the top-
pling of any other species. Succession in this case is a
race for uncontested dominance in recent gaps, rather
thandirectcompetitive interference. N o species is
competmvely superiorto any other. Succession
works on a 'first come, first served' basis - the species
that happen to arrive first become established. It is a
disorderly process, in which any directional changes
are duetoshort-lived species replacinglong-lived
species.
Allogenic and autogenic succession
Autogenic succession is propelled by the members
of a community. In the facilitation model, autogenic
succession is a unidirectlonal sequence of community,
and related ecosystem, changes chat follow the open-
ingup of a new habitat.The sequence of events
takes place even where the physical envlronment I S
unchanglng. Anexample is the heathcycle in Scotland
(Watt 1947). Heather (Calluna vulgaris) is the domi-
nant heathland plant. As a heather plant ages, it loses
its vigour andis invaded by lichens (Cladonia spp.). In
time, the lichen mat dies, leavmg bare ground. The
bare ground is invaded by bearberry (Arctostaphylos
uva-ursi), which in turn is invaded by heather. The
cycle takes about twenty to thirty years.
Allogenicsuccession is driven by fluctuations
and directional changes in the physical environment.
A host of environmental factors may disturb com-
munities and ecosystems by disrupting the interac-
tions between individuals and species. Whena
stream carries siltinto lake, deposition occurs.
Slowly, the lake may change into amarsh or bog, and
in time the marsh may become dry land.
Primary succession
Clements distinguished between prlmary succession
and secondary succession. Primary succession
occurs on newly uncovered bare ground that has not
supportedvegetation before. New oceanic islands,
ablation zones in front of glaciers, developing sand
dunes, fresh river alluvium, newly exposed rock
produced by faulting or volcanic activity, and such
human-made features as spoil heaps areallopen to
first-time colonnation. The full sequence of commu-
nitiesform a primarysere or prisere. Different
priseresoccur on different substrates:ahydrosere
is the colonization of open water; a halosere IS the
colonization of salt marshes; psammosere
a is
the colonization on sand dunes; and a lithosereis the
colonizatlon of bare rock
A hydrosere is recognized in the fenlands of
England (e.g. Tansley 1939). Open wateris colonized
by aquaticmacrophytes,andthen by reeds and
bulrushes. Decaying organic matter from these plants
accumulates to create a reed swamp In which water
level is shallower. Marsh and fen plants become
established in the shallower water. Further accumu-
lation of soil leads to even shallower water. Such
shrubs as alder then invade to produce 'carr' (a scrub
or woodland vegetation),andultimately, so the
classic interpretation claimed, carr would change into
mesic oak woodland.
Glacier Bay, Alaska
A classic study of primary succession was made In
Glacier Bay National Park, south-east Alaska (Cooper
1923,1931,1939; Crocker and Major 1955).The
glacier has retreatedconslderably over the last 240
years (Figure 7.2; Plate7.1).The pioneer stage is
characterized by roadside rock moss (Racomitrium
canesrens) and hoary rock moss (Racomitriunz fanngi-
nosum), broad-leavedwillow herb or river beauty
(Epilobrum latijofium), northern scouring rush
yellow mountain avens (Dryns
(Equisetr~tn variegatum),
drunzmondii), and the Arctic willow (Salix arctica). In
the next stage,the Barclay willow (Salixbarcfayi),
Sitka willow (Salix srtchensis), and feltleafwillow
(Safixalaxensis), undergreen willow (Salix coommutata),
andother willows appear.Theystart as prostrate
forms but eventually develop an erect habit, formlng
dense scrub. Later stages of succession vary from place
 C O M M U NCI T
HYA N G E 175

-
0 10

Figure 7.2 Glacier Bay, Alaska, showing the positions of the glacier terrninl and Fastle's (1995) study sites (referred to
on p. 176).
Source: After Crocker and Major (1955) and Fastie (1995)
176 C O M M U NCI THYA N G E

Plate 7.1 Plant successionat Glacier B ay,Alaska (a) Upper Muir inlet (Site 1 in Fi ure 7.2), 20
old: glacial till surface with young plants of yellow mountain avens (Dryas cf!mmmondii],
cottonwood (Populus trichocorpa) and Sitka spruce (Salix sitrhensis) (b)Goose Cove (Site 3 in Figure
7.2), 60 years old: alder thicket overto ped by black cottonwood (c) Muir Point (Site 6 in Figure
7.2), 105-1 10 years old: alder and wilkw thicket overtopped by scattered Sitka spruce (d) York
Creek (Site 9 in Figure 7.2), 165 years old: Sitka spruce forest (with western hemlock) that was not
preceded by a dense alder thicket or cottonwood forest
Photographs by Christopher L. Fastie

to place. They involve three maln changesIn vegeta- mated by alderthickets.Thlrd,westernhemlock

tion.First,green or mountainalder (Alnus crzspa)
establishes itself In some areas. On the east side of
Glacier Bay (Muir Inlet) and withm 50 years, it forms
almost pure thickets, some 10 m tall, with scattered
mdividuals of black cottonwood (Populus trzrhocarpu).
Second, Sitka spruce(Pzcea srtrhenszs) Invades and after
some 120 years forms dense, pure stands. The spruce
finds I t difficult to establish itself in those areas dom-
(Tsugaheterophylla) enters the communlty, atrivmg
soon after the spruce. Eventually, after a further 80
years or so, spruce-hemlock forest IS the climax vege-
tatlon, atleast on well-drarned slopes. In wetter sites,
where the ground is gently slopmg or flat, Sphagnum
speclesinvade the forest floor. The more Sphagnum
there IS, the wetter the conditlonsbecome. Trees start
to die and wetland forms. The wetlandIS dommated
 COMMUNITY
CHANGE 177

by Sphagnum, with occasional shore pine lndividuals Rakata,Sertung,andPanjangweresterilizedin

1883. Anak Krakatau emerged in 1930 and has a
(Pinus contorta), which are tolerantof the wet habitat.
disturbed hlstory of colonization.
Community succession on each of the three main
Krakatau Islands, Indonesia
islands follows a slmilar patternfor the first 50 years.
The eruptions of 20 May to 27 August 1883 largely The coastalcommunltieswereestablishedrapidly.
or completelysterilizedandgreatlyreshaped the Typical dominant strand-line species were among the
Islands in the Krakatau group. There are four Islands first to colonize, producing a one- or two-phase suc-
in the group - Rakata, Sertung, Panjang, and Anak cession. In the interlor lowlands and upland Rakata,
Krakatau.TheylieintheSundaStraitroughly an early phase of highlydisperslveferns,grasses
equidistant 44 km from ‘mainland’ Java and Sumatra (carrledbywinds),anda few Composltaeslowly
(Figure7.3).Twolarge‘steppingstone’islands - diversified. By 1897they had gainedsuchspecies
Sebesi and Sebuku - connect them with Sumatra. as the heliophilous(sun-loving)terrestrialorchids

.
aurlua

Strait .

Krakatau

- i
fbkata

0 km 5 ‘
0 km 50

Figure 7.3 The location of the KrakatauIsland group. Rakata,Panjang,andSertungareremnants of the pre-1883
volcano; Anak Krakatau, whlch appeared in 1930, is the only volcanlcally actme Island.
Source: After Whlttaker and Bush (1993)
178 COMMUNITY CHANGE

Arundina
graminifolia, Phailrs tankervilliae, and Much of the pattern reflects the differinglife histories
Spathoglottis plicata. A scattering of shrubs and trees, and time to maturityof the component species.
most of which were anlmal-dispersed colonlsts that
were interspersed amongthe savannah vegetation,
soon spread. Forest covered Rakata by the end of the Ghost towns in the western Great Basin
1920s. The forest closure led to the loss of a number TerrillandWonder,which arein central-western
of earlycolonists that were open-habltat species. Nevada,UnitedStates, are abandonedgold-and
Additionally, open areas werereduced to relatively silver-mining camps. Terrillwas abandoned in about
small gaps and fauna dependent on open areas suf- I 7 1 5 and Wonder in about 1925. Secondary succes-
fered. Losses includedbirds such as the red-vented sion followingabandonment was studied for two
bulbul (Py~xonotwtrrfer) andthelong-tailedshrike levels of disturbance - greatly disturbed (abandoned
(Laniw schacb bcntet). roads) and moderately disturbed (wlthin5 m of build-
ing foundation edges) - and at ‘undisturbed’ control
Secondary succession plots (Knapp 1992).Twenty-seven species were found
atTerrill.Cheatgrass (called drooping brome-grass
Secondary succession occurs on severely disturbed
in the United Kingdom)(Bromw tectorum),Shockley’s
groundthat previously supportedvegetation. Fire,
desertthorn (Lyciutnshockleyi), indian ricegrass
flood, forest clearance, the removal of grazing
(Oryzopsis hytnenoides), and Bailey’s greasewood
animals, hurricanes, and many other factors may
(Sarcobatus baileyi) dominatedtheabandoned road
inaugurate secondary succession.
(Plate 7.2a). Shadscale (Atrzplexconfrtifolia), cheat-
grass, and Halogeton glonreratus (an introduced Asiatlc
Abandoned fields in Minnesota annual) dominated the foundation edges (Plate 7.2b).
Before the 1880s, upland habitats in the Cedar Creek The control plotwas dominated by greasewood, shad-
Natural History Area, Minnesota, were a mosaic of scale, and cheatgrass. At Wonder, just seven species
oak savannah (open oak woodland), pralrie openings, were found. Big sagebrush (Arternisla trzdentata) was
andscatteredstands of oak forest, plne forest, and thedominant species intheabandoned road and
maple forest (Tilman 1788). Farmmg converted some control site (Plate7 . 2 ~ )Big
. sagebrush andcheatgrass
of the land to fields. Some fields were abandoned at comprlsed 90 per cent of the cover around foundation
different times andsecondary succession started. Aset edges (Plate 7.2d).
of twenty-twoabandoned fields, placedin chrono- In both towns, a striking feature ofsuccession is the
logical order, provides a picture of the successional higher percentages of therophytes around foundation
process (Figure 7.4).
Theinitial
dominants are peripheries. This is seen in the
highpercentage
annuals and short-lived perennials, many of which, of cheatgrass at Terrill and Wonder, and in the high
including ragweed (Ambrosza artmisiifolia), are agri- percentages of Halogeton and tumbleweed(Salsola kali)
cultural weeds. These are replaced by a sequence of at Terrill (Plate 7.2). They are not so common on
perennial grasses. After fifty years, thedominant highly disturbed abandoned roads because soil bulk
grasses are the little
bluestem (Schizachyrzum density is higher there, which hinders the growth of
sroparirrm) and big bluestem (Andropogon gerardi),both tumbleweed’s andcheatgrass’s extensive rootsystems,
natlve prairie species. Woody plants - mainly shrubs, and because phosphorus and nltrogen levels are low,
vines, and seedlings or saplings of white oak (Quercus whichhamperstheestablishment of cheatgrass.
alba), red oak (Quercus rubra), and white pme (Pinw Vegetation has reverted tosomethingapproaching
strobus) - slowly increase in abundance.After sixty its original state(salt desert shrub in the case ofTerrill
years, they account for about 12 per cent of the cover. and sagebrush-grass in thecase of Wonder), but com-
Many of the successional changes are explained by an plete recovery to a ‘climax’ state is unlikely. Thero-
increase of soil nitrogen over the sixty-year period. phytes, especially cheatgrass,arelikely topersist.
 C O M M U NCI H
TYANGE 179

species. The secondary succession has produced a new

25 1 'climax' vegetatlon, a sort of permanent disclimax.

E G O C E N T R I CN A T U R E :
DISEQUILIBRIUMCOMMUNITIES

IU - The idea that communlties and ecosystems mlght not

be in equilibrium, at leastnot In Clements's sense
of stableclimaxcommunities o r Odum's sense of
balanced ecosystems, was first ralsed by Henry Allan
Gleason and Alexander StuartWatt in the 1920s. The
disequilibrium view rose to stardom in the early
-0
lc)70s, whensomeecologlstsdared to suggest that
in -
c Ticklegrass
3
n I "

succession leadsnowhere inparticular, thatthere

Brambles
are no long-lasting climatlc climaxes (e.g. Drury and
Nisbet 1973). Instead, each species'does ~ t sown
thing',communltiesareever-changing,temporary
0
alliances of Individuals, andsuccession runs in several
directlons. Thls disequilibrium view emphasizes the
Goldenrod
5 - Bush clover indivldualistlcbehaviour of specles andtheevolu-
tionary nature of Communities. It stresses imbalance,
0 I I I I I I
disharmony, disturbance,
and
unpredictability in
0 20 40 60 Nature.And it focuses onthegeographyof eco-
Successlonal age (years) systems - landscape patches, corrldors, and matrixes
replace theclimaxformationandecosystem,and
F i p r r 7.4 Secondary succession In CedarCreekNatural
HistoryArea,Minnesota,United Srates. The diagram I S landscape mosaics replace the assumed homogeneous
basedon an oldfieldchronosequenceandobservations climaxes and ecosystems.
within some permanent plots. The species are quackgrass The indivldualistic comlngs and goingsof species
(Agropyron repem), ragweed (Avtlirosm urtemisiifolia), hlg Influence communitychangein aprofound way.
bluestem (Anrivopogon gerurdi), Kentuckybluegrass (Pua Communities change because new species arrive and
praterrm), little bluestern (Schrzuchyirrm scoparirrm); tlckle-
grass (Agrostrs J L - u ~ ~hoary
u ) , alyssum (Berteroa incuna), old species are lost. New species appear in speciation
hawksheard (Crep" tectorm), horseweed ( E r r p m canaden.rrs), events and through immigration. Old species vanish
red sorrel (Rumex ucetorrl/a), brambles ( R 1 h s spp.), roses through local extinction(extirpatlon)andthrough
(Rosa spp.); wormwood (Arterrma ltrdovtcrana),pennyroyal emigration. Some species increase in abundance and
(Hedeotm hzspzda), bush clover (Lespedeza capitafa), golden-
others decrease in abundance, thus tipping the com-
rod (So/idup netmrulis), Indian grass (Sorghastvm nutans).
Sorwce: After Tilman (1988) petitivebalancewithin a community. Each species
has itsownpropensity for dispersal,invasion,and
Cheatgrassexcelsindisturbedareas. It establishes populationexpansion. Community assembly is an
itself easily ondisturbedsitesandcreates a self- unceasing process of speciesarrivals,persistence,
enhancing cycle by promotingtheoccurrenceof increase,decrease, andextinctions played out in an
additional disturbances - fire intensity and frequency individualistic way.
Evidence
that
communities
and grazing by smallmammals. So, cheatgrass, assemble(anddisassemble) inthismanner is seen
which is a Mediterranean Introduction (p. 172), will in multidirectional succession, as revealed in recent
probably staydominant at the expense ofother, natlve, vegetation chronosequences,and
community
in
180 COMMUNITY CHANGE

Mato 7.2 Secondary succession in Great Basin ghosttowns, United States (a) Terrill- view looking
west. The 'main street' on the rightis barely visible. It leads to the sole remaining skucture. In the
distance is the Terrill water tower. Thesmall,pale,bushy plant is indian ricegrass (Oryropsis
hymenoider). The dominant shrubs are Baile 's greasewood (Sarcobotus buileyi) and Shockley's
desert thorn (Lycium shockleyi) (b) A view of t i e sole remaining building. Speciesin the foreground
are Halogeton glomeratus (mainly in the lower left corner) and tumbleweed (Salsola kali) (next to
the scrap wood) (c) Wonder - view looking north-west towards tailings pile. Almost the entire
area in the foreground was part of the town. The shrubs are near1 all big sagebrush (Artemisiu
tridentarb); the grass is cheatgrass (Bromus tectorum) (d) View of t i e foundation at Wonder. Big
sagebrush, cheatgrass and pition pine (Pinus edulis) arein the foreground
Photographs by Paul A. Knapp
 COMMUNITY CHANGE 181

impermanence, as revealed inpalaeobotanical and based on tree-ring records from 850 trees at ten sites
palaeozoological studies. of different age (Fastie 1995). The findings suggested
that succession there is multidirectional.Thethree
oldest sites were deglaciated before 1840. They differ
Multidirectional succession
from all younger sites in three ways. First, they were
A result of individualistlc communityassembly is that all invadedearly by Sitkaspruce (Plrea SltchenJIJ).
succession may continue along many pathways, and IS Second, they all support westernhemlock (Tsuga
not necessarily fenced into a singlepredetermlned heterophylla).Third, they all appear to have had early
path. Some field studies support this Idea. shrub thickets. Sitka alder (Ahus sinctata) is a nitro-
gen-fixing shrub.Onlyatsitesdeglaciated since
1840 has it been an important and long-lived species.
Hawaiian montane rain forest
Black cottonwood ( P O ~ I Ltrichorarpa)
~UJ has domlnated
Montane rain forest on a windward slope of Mauna the overstoreyonly atsltesdeglaclated slnce 1900.
Loa, Hawaii, displaysa primary successional sequence The new reconstruction of vegetation succession In
on lava flows with ages ranging from 8 years to 9,000 the Glacier Bay area suggests additions or replace-
years (Kitayama et al. 1995). Both downy (pubescent) ments of slngle specles. These smgle-specles changes
andsmooth(glabrous) varletles of thetree Metro- distinguishthree successlonal pathways that occur
JideroJ polytrrorpha (Myrtaceae) dominated the upper in different places andat different tlmes.This re-
canopy layers on all lava flows in the age range 50 to evaluation of the evidencedispels the Idea that
1,400 years. Thedowny varlety was replaced by communities of different ageatGlacier Bay form
thesmooth varietyon the flows morethan3,000 a single chronosequence describingunidirectional
years old. Lower forest layers are dominated a matted successlon.
fern, Dicranoptwis /inearzs, for the first 300 years, The multiple successlonal pathways seem to result
and then by tree ferns (Cibotlctm spp.). The Cibotlutn from the number and timmg of woody species arrlv-
cover declined slightly after 3,000 years, while other ing on the deglacrated surfaces. They do not appear
native herb and shrub species increased. A‘climax’ to stem from spatial differences in substrate texture
vegetatlonstate was not reached - blomass and and lithology. For example, site proximity to a seed
species compositionchangedcontinuouslyduring source at the time of deglaciationaccounts for up
succession. Such divergent succession may be unique to 58 per cent of the variance in early Sitka spruce
to Hawaii, where the flora is naturally impoverished recrultment.
Nitrogen-fixing is another factor
anddisharmonicdue to Its geographlcal lsolatlon influencing successional pathways.Long-livedalder
(Kitayama et al. 1995). Against thls vlew, it could be thicketsproduce soil nltrogen pools thattendto
argued that Hawaii is the sort of place where succes- reduceconifer recrultmentand prevent the rapld
sion should have the bestchance of following the development of spruce-hemlock forest.
classical model (R. J. Whlttaker pers. comm.). It is
very interestmg that montane forest on Hawaii does
Krakatau Islands revisited
not follow the classical model. Rather, it may be an
example of a general pattern of non-equilibrium The latest studies of prlmary succession on Krakatau
dynamics of thekind revealed by new work at favour adisequilibriumlnterpretatlon(e.g. R . J.
Glacler Bay and the Krakatau Islands. Whlttaker et al. 1989, 1992; Bush et al. 1992; R. J.
WhittakerandJones1994).Withthe exceptlon
of thestrand-line specles, all components of the
Glacier Bay revisited
Krakatau fauna and flora are still changlng. A lastlng
A re-evaluatlon of the succession In Glacler Bay, equilibrium does not prevail because the faunal
Alaska, used reconstructlons of standdevelopment and floral diversity I S contlnuallybeingaltered by
182 COMMUNITY CHANGE
dispersal opportunities,and by disturbance events
ranging from continuing volcanism tothe fall of
individual trees. The patchy nature of these processes
helps to explainwhy the forest-canopy architecture
on PanjangandSertungchanged materiallyfrom
1983 to 1989, while the pace of change on Rakata
over the same period was fairly slow. It also accounts
for slgnificant differences within and between forests
on the four islands. Overall, the forest dynamlcs of
Krakatau have been highly episodic, withstands
experiencmg times of relatively minor change punc-
tuated by pulses of rapid turnover and change; andall
the while, new colonists species have arrived and have
spread (R. J. Whittaker pers. comm.).
O n Rakata, for example,there arefour common
successional pathways that have produced distinct
forest communities (R. J . Whittaker and Bush L993).
The first occurs on coastal communities, where either
Tenninalia r.atuppa-Batrir/gtonta a.rluti1.u woodland I S
rapldly established, or else the same woodland
is established after a stage of Cmuarrnu equiset$olia
woodland (Plate 7.34. Path two is followed inland
(butnotinuplands)andruns fromferns, to grass
savannah, to Macaranga tanarrus-Ficus spp. forest. to
Neonauclea calycina forest (Plate 7.3b). Path three
the same as path two, but the current stage is Ficus
calycina forest (Plate 7 . 3 ~ )Path
prrh1nervz~-Neonautlea4~.lea
four occurs in the upland (Plate 7.3d). It runs
ferns, to grasssavannah, to Cyrtandra sulcata scrub,
to SrheffIera polyhotrya-Sar/ralria nudipora-Ficus ribes
submontane forest.
The vegetation of Rakata is quite differentfrom
thevegetationonPanjangandSergung(Tagawa
1992). Neonauclea calycina seeds successfully formed
forest on Rakata, but failed to do so on Panjang and
Sertung. Factors in seed dispersal, soil conditions, and
disturbance by volcanic activity on Anak Krakatau all
help to explain these differences. The Tirnonius corn-
pressicaulis and Dysoxylum gaudichaudianunr forests
foundonPanjangandSertung developedon both
islands after the appearance of Anak Krakatau. Both
Timonius romnpressiraults andparticularly Dysoxylurn
gaudilhaudianum have larger seeds than Neonauclea
and they were able to germinate and grow under the
regeneratedmixed forest canopy, but it was not so
easy for themto colonize the Neonauclea forest.
Dysoxylrm ga~rdichatrdianurninvasion progresses only
gradually on Rakata.
Fleeting communities
Communities,likeindividuals, are impermanent.
Species abundances anddistributions
constantly
change, each according to Its own life-history charac-
teristics, largelyin response toanever-changing
environment.This communityimpermanence is
seen in thechangingdistributions of threesmall
North American mammals since the Late Quaternary
(Figure7.5).Thenorthernplains pocket gopher
(Thornomnystalporde.r) lived in south-western Wisconsin
around17,000 years ago and persisted in western
Iowa until at least 14,800 years ago. Climatic change
associated wlth deglaclation then caused it to move
west. The samc climatic change prompted the least
shrew (Ctypto/i.rpatva) to shift eastwards. The collared
lemming (Dimstonyx spp.), which lived in a broad
band south of the LaurentldeIce sheet, went 1,600 km
to the north.
With each species actingindividually, it follows
is that communities, both local ones and biomes, will
comeandgo in answer toenvironmentalchanges
. (e.g.GrahamandGrimm1990).Thisargument
from leads toamomentous conclusion: there is nothing
special about present-day communities and biomes.
However,this boldassertion should be tempered
with a cautionary note - insect species and commu-
nities have shown remarkable constancy in the face of
Quaternary climatic fluctuatlons (Coope 1994).
Communityimpermanence is evidencedin three
aspects of communities - in communities with no
modernanalogues, in communitiesthat are dis-
harmonious,and in communities that behave
chaotically.
No-modern-analogue communities
Some modern communities and biomes are similar to
past ones, but most have no exact fossil counterparts.
Contrariwise, many fossil communitiesand biomes
have no precise modern analogues. Thisfinding
 COMMUNITY CHANGE 183

Plate 7.3 Successional pathways on Rakata (a) Pathone:Casuarinauiseh’foliawoodland (at

land Ficusubinervis-Neonaucleacalycinaforest (d) Path four: upland forest

7
north end of AnakKrakatau) (b) Path two: lowland Neonauclea calycina orest (c) Path three: low-

Photographsy ! R. J. Whittaker

supportsthedisequilibrium view of communities. nificant plant community components (Delcourt and

The list of past communlties that lack modern ana-
l o p e s is growing fast. In the Missouri-Arkansas bor-
der region, United States,from 13,000 to8,000 years
ago, the eastern hornbeam (0.rtyra vtrginiana) and the
American hornbeam (Carpinla carolintana) were sig-
Delcourt1994).Thesecommunlties,which were
foundbetween the Appalachian Mountains and the
OzarkHighlands, bore little resemblance to any
modern communitiesIn eastern North America. They
appear to have evolved In a climate characterized by
184 COMMUNITY CHANGE

(a) Northern
plains (b) least shrew (c) Collared
lemming
gopher pocket Cryptotis parva Dicrostonyx spp.
Thornornys talpordes

Late Pleistocenefossillocalities of taxa - Southernlimit of Wisconsinglaciation

Figure 7.5 Individualistlc response of some small North Amerlcan mammals since the LateQuaternary.(a) Northern
plalns pocket gopher (Thmmys tulpozder). (b) Least shrew (Ctyptotzr purva). (c) Collared lemming (Dzmstonyx spp.).
Source: After Graham (1992)

heightened seasonality and springtime peaks in solar 1987).InthesouthernGreatPlainsandTexas,

radiation. Farther north, in the north-central United UnitedStates,present-daygrassland or deciduous
States, a communlty richIn spruce and sedges existed forest species - Including the least shrew (Cvptotrs
from about 18,000 to 12,000 years ago. This commu- pawa), theboglemmlng (Synapotomyscoopwi), the
nity was aborealgrasslandbiome(Rhodes 1984). pralrievole (Mimotus ochrogastw), andshort-tailed
It occupied a broad swath of land south of the ice shrews (Blarina spp.) - livedcheek-by-jowlwith
sheet and has no modern counterpart, though it bore present-day boreal specles- including the long-tailed
some resemblance to the vegetatlon now found in theshrew (Swex cznweus), white-tailed lack rabbit ( k p u ~
southern part of the Ungava Penlnsula, in northern townsendii), ermlne or stoat (Mustefa errnmnea), and
Quebec, Canada. meadow vole (Microtus pennsyfvanrcus) (Lundelius et
al. 1983; also see Figure7.5).DuringtheLate
Pleistocene epoch, disharmonious animal communi-
Disharmonious communities
ties were found over all the United States, except for
The fauna and flora of communltles with no modern the far west where vertebrate faunas bore a strong
analoguesarecommonlydescribed as disharmo- resemblancetotheirmoderndayequivalents,and
nious communities. This inapt name Inadvertently date from at least 400,000years ago to the Holocene
conjures an Imageof animal and plants struggling for epoch.Thesedisharmoniouscommunities evolved
survival in an alien environment. It IS meant to con- from species respondingindividually tochanging
vey the idea that these communltles had evolved in, environmentalconditlonsdurlngLatePleistocene
and flourished under, climatic types that no longer times (Graham 1979). At the end of the Pleistocene,
existanywhereintheworld(GrahamandMead new environmental changes led to the disassembly of

the communitles. The climate became more seasonal
andindividual species had to readjust thelrdis-
tributlons. Communltles of a distinctly modern mark
emerged during the Holocene epoch.
North America does not have a monopoly in
disharmonlouscommunlties.InAustralia, an Early
Pliocenefauna from Victoria - the Hamilton local
fauna - contains several extant genera whose livmg
species live almost exclusively in rain forest or rain-
forest frlnges (Flannery et al. 1992). The lndicatlon
is, therefore, thatthe Pliocene fauna lived In rain-
forest environment, but a morecomplex raln forest
than exlsts today. Modernrepresentatives of four
genera (Hypsip’ytunodon, Dorc.opsir, Detldrdagm, and
Strrgomscxs) are almostentlrely rain-forest dwellers,
but they live in different kinds of r a n forest. Livmg
specles of D0rrop.rz.r (kinds of kangaroo) live In high
mountam forests, lowland ralnforests, mossy montane
forests, andmid-montane forests. Livlngspecies of
Dendrofugus (tree kangaroos) live mainly in montane
rain forests. Hypsiprymnodon mrtssthatr/.r (themusky
rat-kangaroo) is restrlcted to rain forest where it
prefers wetter areas. The modern New Guineaspecies
Strigocxsws gytmotns is chiefly aran-forestdweller,
though it also lives In areas of regrowth, mangrove
swamps, and woodland savannah. Livlng species
of ThyLogale (pademelons) live In an array of environ-
ments, including rain forests, wet sclerophyll forests,
andhighmontane forests. Othermodern relatives
of the fossils In the Hamilton fauna, which includes
pseudochelrids, petaurids, and kangaroos and walla-
bies,livein awide range of habltats.Itcontains
two species, Tr/cho.rur//.I(brushtail possums) andStrrgo-
c m x r (cuscuses), whose ranges do not overlap at
present. I t I S thus a disharmoniousassemblage. Taken
as a whole, theHamiltonmammalian assemblage
suggests a diversity of habitats in the Early Pliocene.
The environmentalmosaic consisted of patches of rain
forest, patches of other wetforests, and open area
patches. Nothing like thisenvironment is known today.
Chaotic communities
Inthe 1990s, the notion of non-equilibrium was
formalized In thetheory of chaoticdynamics.
COMMUNITY CHANGE 185
The main idea IS that all Nature, including the com-
munitiesand ecosystems, is fundamentallyerratic,
discontinuous,andinherentlyunpredictable.This
V I ~ W of communities and ecosystems arose from
mathematical models. Themodels confirmed what
ecologists had felt for a long time buthad been unable
to prove - emergentcommunltyproperties, such
as food webs anda resistance to invasion by alien
species, arise from the host of indivldual interactions
In an assembling community. In turn, the emergent
community propertles influence the local interactions.
All evolving ecosystems, from the smallest pond to
the entlre ecosphere, possess emergent properties and
appear to behave like superogan~sms. But this super-
organic behaviour I S the result of a continuing two-
way feedback between local interactlonsandglobal
propertles. It is not the outcome of some mystical
global property determmng the local interactlons of
system components, as vitalists would contend. Nor
is it the cumulatlve result of local mteractlons in the
system, as mechanists would hold.No,the whole
system is an Integrated, dynamic structure powered
by energy and involvmg two-way Interaction
between all levels.
Experiments wlth ‘computer communitles’showed
that species-poor communmes were easy to invade
(Pimm 1991). Communmes of up to about twelve
species offered essentlally open access to intruding
specles. Beyond that number, in specles-rich commu-
nlties, there were two results. First, newly established
species-rlch communltles were more difficult to
Invade than specles-poor communltles. Second, long-
established communitles were even harder to Invade
than newly established species-rich communities.
Other mathematical experiments started with a 125-
species pool of plants, herbivores,carnivores, and
omnivores(Drake 1990). Specles were selected one
at a time to join an assembling community. Second
chances were allowed for first-time failed entrants. An
extremely persistent community emerged comprising
about fifteen species. When the model was rerun with
the
same species pool, an extremely perslstent
communlty again emerged, but this time withdiffer-
ent component specles than in the first community.
There was nothing speclal about the specles In the
186 C O M M U N I T YC H A N G E

communities: most species could become a member dissected by roads, brokenintodiscrete patches
of a persistent community under the r ~ g h tcircum- by felling, and the newly created patches are shrink-
stances; the actual species present
depended on ing and someof them disappearing through attrition.
happenstance. It was the dynamics of the persistent The overall effect of land-cover change is habitat
communities that was special: a persistent commu- fragmentation.
nity of fifteen species could not be reassembled from
scratch using only those fifteen species. This finding
Habitat fragmentation
suggeststhatcommunitiescannot be artificially
manufactured from a partlcular set of species - they This is thebreakingup of large habitats or areas
have to evolve and to create themselves out a large into smaller parcels. It is enormously significant for
number of possible species interactions. wildlife and poses a global environmental problem.
As habitat patches become smaller and more isolated,
so several community changes occur. The numbers of
UNDER THE AXE AND PLOUGH: generalist specles, species that can live in more than
LAND COVER TRANSFORMATION one habltat, edge species, and cxotlc specles all rise.
The nest predation rate increases, populations fall,
Inalmost allparts of the world, biomesare being and extlnctions become more common. The numbers
changed on a masswe scale. Much of this land-cover of species specialized for life within a habitat intertor,
transformation has taken placein the past two and species with a large home range, both decrease,
centuries (Buringh and Dudal 1987) (Figure 7.6). It as does thediversity of habitatinterior species.
resulted from the western European core region Habitat
fragmentation also changes ecosystem
pushingout successive waves of exploltationinto processes. It disturbs the integrity of drainage net-
peripheral regions of theglobe. As thefrontiers works, affects water quality in aquifers, altersthe
expanded, so isolated subsistence economies were disturbanceregimeto which species have adapted,
drawn irresistibly into a single world market. Basic and influences other ecosystem processes.
commodities - crops, minerals, wood, water,and The effects of fragmentinghabitats onwildlife
wild animals - were traded globally. Combined with are evident in the examples of the skipper butterfly
steam power, medicine, and advances in agricultural in Britain,the malleefowlin Australia,andthe
technology,theseeconomic changes set in motion reticulated velvet gecko in Australia.
the 'great transformation' - a world-wide alteration
of land cover.
The skipper butterfly in Britain
The land-cover transformation has changed
biomes.From 1860to1978,8,517,000 km' of In Britain, the skipper butterfly(Hespevra comma) lives
land were converted to cropping (Revelle 1984). The in heavily grazed calcareous grasslands (Colour plate
converted land was originally forest (28.5 per cent), 11) (C. D. Thomas and Jones 1993). The grazing IS
woodland (18.3 per cent), savannah (13.8 per carried out largely by rabbits (0ryrtohpr.r cwrnicdrrs).
cent), othergrassland (34.9 per cent), wetland (2.5per Durtng the mid-l95Os, the rabbits were devastated
cent),anddesert(2.0 per cent).Amajorproblem by myxomatosis and the habltat became overgrown.
with thls ecosystem conversion is itspatchynature The skipper butterfly populatlon shrank and became
- a field here, a clearlng there - that has broken metapopulation,
a occupying forty-six or fewer
the original habitat into Increasingly Isolated habltat localitles in ten refuges (Figure 7.7). The rabblts had
fragments. Land-cover change I S caused by five recovered by 1982, and many former grassland sites,
processes - perforation, dissection,fragmentation, now neatly cropped, appeared suitable for recolon-
shrinkage,andattrltlon(Forman1995:40&15). izatlon by the skipper butterfly. Indeed, from 1982
Forests, for example, are belng perforated by clearings, to 1991,thenumber of populatedhabitat patches
 COMMUNITY CHANGE 187

15

L
a,
0
u
-0
m
J 5

0
900 1100 1300 1500 1700 1900

Figure 7.6 Land-cover transformation, A D 900-1977. The greatest transformation took place In the last two cenrurles.
Source: After M. Williams (1996)

increased by 30 percentin theSouthandNorth The malleefowl in Australia

Downs, with most of the increase taking place in East
Sussex. Most of the recolonized habitat patches were
fairly large and close to other populated patches. But
even by 1991, many suitable habitat patches had not
been reoccupled. Were the habitat not fragmented,
theskipperbutterflymight be expected to fill its
previousrangein south-eastEnglandwithln fifty
to seventy-five years. Inthefragmented landscape,
it is likely that little or no further spread will take
place inthe twenty-first century, except In East
Sussex. In most places, bands of unsultable habitat,
whlcharemore than 10 km wide, haltthe spryad.
Conservation of theskipper butterflyrequires the
protection of metapopulations in networks of habitat
patches.
Mallee is an Australian sclerophyllous shrub forma-
tion (equivalent to the maquis in the Mediterranean
region)characterized by high bushes of shrubs and
small trees. Loss, fragmentation, and degradation of
mallee habitat within the New South Wales wheat-
belt have caused a marked decline in the range and
local abundance of malleefowl (Lezpoa o d a t a )
(PriddelandWheeler 1994). The malleefowl is a
large, ground-dwelling bird that makes a mound for
its nest and keeps the temperature tightly controlled.
Small,disjunct
populations of malleefowl now
occupy small and Isolated remnants of mallee habi-
tat. Several of these populations have recently become
locally extinct. As an experiment, young malleefowl
188 COMMUNITY CHANGE

..... Pre-1920
range

................. 192061 range

0 Refuge distribution, 1982

1982 and 1991 distribution,

' excludingpost-1975recolonization
Recolonization since 1975, occupied in 1991
Suitable habitat unoccupied in 1991

Figure 7.7 Skipper butterfly (Haperzu ronmu) decline in England durlng the twentleth century The three published maps
show the sklpper butterfly beyond the range shown, but each of the five records occurs on just one of the maps and they
are excluded. The bottom map shows the results of a recent survey.
Soutre: After C. D. Thomas and Jones (1993)
 COMMUNITY C H A N G E 189

(8-184 days o l d ) reared in captivity were released in treespecies composltion,andvegetationstructure.

March and June1988 Into a 55-ha remnant ofmallee ord/(ru popuiatlon sizes vary consldetably among

vegetation that contained a small but declinlng pop- remnants.Theyarepoorlycorrelatedwithremnant

ulation of malleefowl. From the first day after release, area orthenumber of smooth-barkedeucalypts.
malleefowl were founddead.
Deathscontlnued Population slze does notappear to be limltecl by
until, wrthrn a relatively short tlme, no malleefowl habltat availability In most remnants. The number of
remainedalive. Themain cause of theirdemlse adults of breeding age is small In most populatmns
was predation, which accounted for 94 per cent of the suggesting that they may be susceptible to stochastic
deaths.Raptorsaccounted for 26-39 percent o f extlnctlon pressures. The specles I S a poor clisperser
the loss, andintroducedpredators, chiefly the red - interactionbetween local populations is small or
fox (V~/peslw/p<,.r),accounted for 55-68 percent. zero, even over distances of 700 m or less. Thls
Helpings of supplementary food made no difference poor dispersalability means thatthepossibility of
to survival. Young malleefowl rely principally on remnantrecolonizatlonafteranextinctionevent IS
camouflage for safety. They have no effectlve defence unlikely. The occupancyrate of Ord/wcr rctl(-/(/L(tL(
or escape belmvlour to evade ground-dwelling preda- In remnant woodland is thus likely to clecline, and
tors. Foxes are ~rnposlngsevere predation pressure the species is likely to becomerestrictedto a few,
onyoungmalleefowl,andareprobablycurtailing large remnants. It may therefore be futile to clirect
recrultment into the breeding population. Foxes are conservationeffortIntoprotectingsmallwoodland
thus a m a p r threat to the continuance of malleefowl remnants of a few hundred hectares.
remnant populations in the
New South
Wales
wheat-belt.
Loss of wetlands
The world's wetlands are theswamps,marshes,
An Australian gecko
bogs,fens,estuarles,saltmarslles, andtldal flats.
The reticulated velvet gecko (Oed/~rrrret/w/utu) is They cover about 6 per cent of the land surface and
restricted In distributlontosouth-westWestern Include s o m e very productive ecosyscems. Butthls
Australia (Sarre 1995) (Colourplate 1La). Withln figure is Falling as wetlands are reclaimed for agricu-
this reglon, it occurs mainly in smooth-barked euca- turdor residential land,
drowned b y clam and
lypt woodland lmbltat, malnly conslstlng of glmlet barrage schemes, or used as rubbish clumps. Sea-level
g u m (Emir/y/>t/ts.rd/{hris)and salmon gum ( E N ( U / ~ / I ~ N .rise
I duringthe twenty-firstcentury also poses a
Much of the woodland has become m a p r threat t o coastal wetlands. The following two
.w/t~/r)ttr~pbioiu).
fragmented by clearing and remalns as Islands wlthin cases illustrate the complexity of changes in wetland
a sea ofwheat (Colour plate 12b).For example, before habitats.
land clearance, about 40 per cent of the Kellerberrin
regmn contamed smooth-barked eucalypt woodland
Bottomland forests in the Santee River
suitable for reticulatedvelvetgeckopopulatlons.
floodplain
The same habltat now occup~es~ 1 s 2t per cent of the
reglon. Only fourremnants,includingthreenature The coastal plain dramage of the Santee River, South
reserves, contalnstands of morethan 570 gimlet Carolina,UnltedStates, was drastically modified
gum and salmon gum trees. Demographic character- in 194 I . Almost 90 percent of theSanteeRlver
istics of nine &dwu populatlonsthatcurrently discharge was diverted into the Cooper River as part
survive In Kellerberrln eucalypt woodland remnants of a hydroelectric power pro~ect (Figure 7 . 8 ) . How-
were assessed and compared with Oec//~r'irpopulatlons ever,thls cliverslonof water has led tosilting In
In three nature reserves. The woodland remnants vary Charleston Harbor, the main navigatlon channels I n
in arm from 0.37 to 5.40 ha, but are slmilar in age, which requlre year-round dredglng. To allcvlate this
190 COMMUNITY CHANGE

0 20 km

Georgetown
VY d

Figure 7.8 Location of the Santee River and proposed river diversion.
Pearlstine et al. (1985)
Sourre: After

costly problem, authorization was granted in 1968 to Aforestgrowthmodel,calledFORFLO, was

redivert most of the water back into the Santee Rlverdevelopedtoquantifytheeffects of thechanged
throughan 18.5 kmlongcanalfeeding off Lake hydrologicalregime
on the
bottomlandforests
Moultrie(Figure 7.8). Anadditionalhydroelectric (Pearlstine et al. 1985) (Box 7.1).I t was used to sim-
power plant wouldbe constructed on this canal.Fears ulate the effect of the proposed river rediversion, and
have been voiced that this rediversion would inun- a modified version of it, along a 25 km reach of the
date much of the Santee floodplain and cause a sub- Santeeforestedfloodplainfrom the rediversion site
stantial decline in the bottomland forest. After the downstreamtoJamestown(Figure 7.8). Inthe
diverslon, the Santee River would return to 80 per proposed rediversion, flow from Lake Marion to the
cent of its pre-diversion flow rate with seasonal spates Santee Rlver staysthe same, flow to the Cooper River
offloods and stages of low flows. A crucial change IS reducedto 85 m3/s(the level whlchprevents
would be a reductlon of flow early in the growing silting in Charleston Harbor), increasing the annual
season. flow to the Santee River vla the rediversion canal to
 COMMUNITY CHANGE 19 1

Box 7.1
KEY FEATURES OF F O R F L O is the duration of the annual flood. Each species has
a tolerance to flooding, and will survive if its range
The FORFLO model allows hydrological variables of tolerance should fall within theflood duration for
to influence tree species composition through seed the plot. Third, the optimum growth of trees was
germination, tree growth, and tree mortality. Key reducedby, amongotherthings,awater-table
features of the model are the assumed relationships functionthatmodelsfloodplainconditions.The
betweenfloodingandvariousaspects of forest water-table
function modifies the tree-growth
growth succession.First,for alltreespecies, save equation to account for the tolerance of species to
black willow (Salix n i p ) and eastern cottonwood the level of water on the plot during the growing
(Populus deltoides),seeds will not germinate when the season. The model computes the height of water
ground is flooded.If the plot should be continu- for each half month during the growing season. It
ously flooded during that period of the year when a is assumed that all trees will fail to grow during the
species would germinate, then the germination of half months when they are more than three-quarters
that species fails. Black willow and eastern cotton- submerged by flood water. At lowerlevels of
wood can germinate whether the land is flooded or submergence, tree growthwas related to water level
not. Second, after having germinated, the survival by a curvilinearfunctioninwhichtheoptimum
of seedlings depends on environmental conditions. water-tabledepth foreachspeciesis takeninto
A notable determinant of the seedling survival rate account.
L

413 m3/s. The modified rediversion was the same as The effects of theproposedrediversionandthe
the proposed rediversion, except that during early the modifiedrediverslonversionon the frequency of
growing season(April toJuly), flow throughthe habitat types are indicated in Figure 7.10. In both
rediversioncanalwouldbekepttoalevelwhich cases there is a large loss of bottomland forest: a 97
could be handled by just one of the three turbinesat per cent loss in the case of the proposed rediversion
the power station. Although this would mean that and a 94 per cent loss in the modified rediversion.
the Cooper River would exceed the critical flowof The saving grace of the modified rediversion plan is
85 m3/s for the four months of the growing season, that a forest cover is maintained bottomland forest
and thus cause some silting at the coast, it might changes to cypress-tupelo forest, rather than to open
promote the preservation of the bottomland forest. water. If these predictions of sweeping changes In
The simulated responses of the forests to the re- the bottomland forest along the Santee River should
diversion are shown in Figures 7.9a and b. The annual be trustworthy, then plainly it would be advisable
duration of flooding is a crucial factor in determining to rethink the plans for rediverting the flow from the
the course of vegetational change. With an annual Cooper River.
flood duration of more than 30-35 per cent (Figure
7.9a),bottomlandhardwoodforestis replaced by
A coastal ecosystem in southern Louisiana
cypress-tupeloforest,baldcypress (Taxodiumdis-
tzchum) and water tupelo (Nyssa uquatzcu) being the Ecosystems occupyingcoastallocationsareunder
only species that would manage to regenerate. When threat from a variety of human activities: gas and oil
annual flood duration was above 65-70 per cent, no exploration, urban growth, sediment diversion, and
specles was able to survive and the forest was replaced greenhouse-induced sea-level rise, to name but afew.
by a non-forest habitat; this happened more rapidly inTo protect and preserve these ecosystems, it is valu-
the subcanopy. able to know what the effects of proposedhuman
192 COMMUNITY CHANGE

-
(b)
Canopy
Bald
cypress Water

/
‘1, tupelo
i
1 Bald

Mockernut hickory

“E
e
“- 0 I
-
t
1 Sub-canopy
I I

lo
1 Sub-canopy

0 60 120 180 0 60 120 180

Time (years)

Fzxrrre 7.9 Results of simulations. (a) Bottomland forest community subjected to an annual flood duration of 45 * 4 per
cent. (b) Bottomland forest community subjected to an annual flood duration of 72 t 5 per cent. The flood duration is
the percentage of a year during which a plot is flooded. The plant species are bald cypress (Taxodiut~rdfJftfbttirJl),
water
tupelo ( N ~ J Jayrrarrfu),
U and sassafras ( S a ~ ~ a f a/brdtm)
va~
Sorwe: After Pearlstine et al. (1985)

activities are likely to be, and how these effects differ scapes in the world. The Atchafalaya River is one of
from natural changes. These questions were addressed thetwoprincipaldistributaries of the Mississlppi
in the Atchafalaya Deltaand adjacent Terrebonne River. It carrles about 30 per cent of the Misslssippi
Parish marshes in southern Loulsiana, Unlted States, dischargetotheGulf of Mexico. Since themid-
using
mathematical
a modelto
simulate likely 1940s, sedimentstransported by the Atchafalaya
changes(Figure 7.11) (Costanza et al. 1990). This River have been lald down in the bay area. In con-
landscape, part of the Mississippi River distributary sequence, the bay area has gradually become filled In,
system, is one of themost rapidly changingland- and in 1973 a new subaerial delta appeared that has
 COMMUNITY CHANGE 193

8.000 1 Bottomland hardwood forest

scenarlos consider what the system would have been
like had not the environment been altered by human
actlon, and if climatic conditions had been different.
Boundary scenarios delve Into the potentla1 impacts
of naturalandhuman-inducedvariationsinthe
boundary conditions of the system, such as sea-level
rise. The management and boundary scenarios were
run by restartlng the model with the actual habitat
map for 1983, rather than the predicted habitat map
for that year, toaddmorerealism.Climateand
historical scenarios were run startlng in 1956,so that
the full impacts of climate and historical varlations
Present After proposed After modified
rediverslon
rediverslon could be assessed. After 1978, the rate of canal and
levee construction for oil and gas exploration slowed
Figure 7.10 Habitat changes in theSantee Rwer study
appreclably compared with the 1956-78 period, so
area.
Sourre: After Pearlstine et al. (1985) all scenarios assumed that no canals nor levees were
built after 1978, except those specifically mentloned
smcegrowntoabout SO km2.Otherchangesare In the scenarlos.
taklngplaceinthe area. The westernTerrebonne It is clear from Table 7.1 that the assumptlons
marshesarebecoming lesssalty,whiletheeastern made about climatic change have a big influence on
part of the area is becoming more salty:the boundary habitatdistributlon by the year 2033.Themean
between fresh and brackish marsheshas shifted closer climate scenario, wherein the long-term average for
to the Gulf in the western marshes, and farther inland each varlablewas used for all weeks,produced a mod-
in theeast (Figure 7.12).As a whole,the study region est loss in land area. On the other hand, the weekly
is losing wetland, but rate of loss in the Terrebonne average climate scenario, wherein each weekly value
marshes has slowed and reversed since the mid-1970s of each climatic variable for the entire run from 1956
owing to m e r deposition. The hydrology of the area to 2033 was set to the average value for that week
has been greatly altered by dredging of waterways in the 1956-83 data, produced a drastic loss of land
andthediggingof access canals for petroleum area. Thisfindingindicatedthattheannual flood
exploration. cycle and other annualcycles In climatic variables are
Adynamlcspatlalmodel was developedand importanttothelandbuilding process, butthat
christened the Coastal Ecological Landscape Spatial chance events, such as major storms and floods, tend
Simulation model(CELSS model for short).It divided to have a net erosional effect on marshland. If the
the marsh-estuary complex Into 2,479 square grid- globalclimateshouldbecome less predictablein
cells, each with an area of 1 km’.The model was the future as a result of global warming, then the
used to predict changes of habitats under a range stability of coastal marshes may be in jeopardy.
of climatic,management,hlstorical,andboundary Several management scenarlos were run. As the data
scenarios. The results of severalscenarloanalyses, In Table 7.1 Indicate, the largest loss of land would
which predicted changes to the year 2033, are sum- arise from the full six-reach levee extension scheme
marized in Table 7.1. Climate scenarios take ‘climate’ that hadbeenconsidered atonetlme.Withthis
to mean all the driving variables including rainfall, scheme, 48 km2 of brackish marsh and 6 km2 of fresh
AtchafalayaRivet flow, wind,and sea-level. They marshwouldbelost by 2033,largely because the
address the impact of climatic changes on the study extended levees prevent sediment-laden water reach-
area. Management scenarios lookat the effect of spe- ing the brackish marsh bordering Four League Bay,
cific human manipulations of the system. Historical wheremost of the loss occurs.Boundaryscenarios
194 COMMUNITY
CHANGE

I I I I
91 w 09w
,31’N

Proposed
freshwater Avoca Island
Palmetto Weir
diversion
Avoca Lake

*
Existing Avoca Swamp forest
Island Levee Freshmarsh
Atchafalaya River Brackishmarsh
Salt marsh
Open water
Upland

Atchafalaya Delta *j Falgout Weir

I” Lake de Cade

Large

Lake’
Lost , l o-mk

Figure 7. I I The Atchafalaya Delta andTerrebonneParishmarshes study area In southernLouisiana, Unlted States.
(a) General location map. (b) Malor geographlcal features, types of habitat In 1983, and management options considered
in the simulations.
. (1990)
Source: After Costanza et a
!
 COMMUNITY
CHANGE 195

1956 p
&\ 1978

...
...
...
1

Swamp forest
Freshmarsh
Brackish marsh
Salt marsh
Open water
Upland

0 50 km

Figwe 7.12 Observed distribution of habitats in the Atchafalaya-Terrebonne study area.

Source: After Costanza et
tal. (1990)

considered the effects of projected rates of sea-level
rise, both high and low projections, on the area. The
results were unexpected. Surprisingly, doubling the
rate of eustatic sea-level rise from 0.23 to 0.46 cmlyr
caused a net gain in land area of 10 km2 relative to
the base case. This was probably because, so long as
sediment loads are high, healthy marshes can keep
pace wlth moderate ratesof sea-level me.
Two historicalscenariosweretested. The first
probed the changes in the system that might have
taken place had not the original Avoca Island levee
been built. The second considered the changes that
might have ensued had not the Avoca levee nor any
of the post-1956 canals been constructed. The results
shown in Table 7.1 suggest that the original levee
and the canals had a major influence on the develop-
ment of the system, causing a far greater lossof land
than would have occurred in their absence.
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GETTINGWARMER:COMMUNITIES
IN T H E T W E N T Y - F I R S T C E N T U R Y
Individuals and communities, by processes of natural
selection, become adaptedtotheenvironmentin
which they live. If that environment should change,
life systems must adapt, move elsewhere, or perish.
Species under pressure
The fate of many species in the twenty-first century,
as the world warms and habitats shrink or vanlsh, is
worrying (Peters 1992a, b). It seems that extinctlon
will continue apace and global biodiversity will drop.
But which species are the most vulnerable to global
warming? Thesafest species are mobile birds, insects,
and mammals who can track their preferred climatic
zone. In the Brltlsh Isles, the white admiral butterfly
(Ludoga ranriiia) and the comma butterfly (Polygonia
c-album) expanded their ranges over the past century
as temperatures rose by about0.5"C (Ford 1982).
However, even mobile specieswould need a food
source, andtheir favourlte menuitemmight be
extinct or have wandered elsewhere.
Five kinds of species appear to be most at risk:
1 Peripheralspecies.Populatlons of anlmals or
plants that are at the contractlng edgeof a species
range are vulnerable to extinction.
2 Geographicallylocalizedspecies. Many cur-
rently
endangered species live
in alarmlngly
limited
habltats.
The
golden lion tamarin
(Leontidexs vosaiicr) I S a small primate that lives in
lowland Atlantic forest, Brazil (Colour plate
13). Lowland Atlantic forest is one of the most
endangered rain forests in theworld. I t once
covered 1 million km' along the Brazilian coast-
line. Now, a mere 7 per cent of the original forest
remains, putting the golden lion tamarin under
enormous threat (but see p. 212).
3 Highly specialized species. Many species have a
close assoclatlon with only one other specles. An
example is the Everglade (or snail)kite (Rostr-
hamus sociabiiis) that feeds exclusively on the large
and colourf~~l Ponzuceu snail inFlorida wetland.
C O M M U NCI H
TYANGE 197
Swamp drainage has already robbed the kite of its
prey in some areas.
4 Poor dispersers. Manytrees have heavy seeds
that are not broadcast far. Plants with limited dis-
persal abilities have knock-on effects alongthe
food chain. Some birds, mammals, and Insects are
closely associated with specific forest trees, which
cannotmigraterapidly.These species wouldbe
hard pressed to survive. TheendangeredKirt-
land's warbler (Dendrozca kirtiandii), for example,
breeds in and nests only upon the ground under
jack-pme (Pinus banksiana) forest on well-dralned
sandy soil in north-central Michigan,
United
States. But notany jack-pine forest will do - it
has tobeyoung, secondary growth forest that
emerges In the aftermath of a forest fire. As the
globalthermometer rises, the jack pines may
move northwards onto less well-dramed soils, and
theKirtland's warbler may find Itself without
any suitablenestingsites(Botkin et ai. 1991).
This specles could be the first casualty of global
warming.
Climatically
5 sensitive
species. Specles in
climatlcally sensitive communities are vulnerable
toglobalwarming.Communities in thls class
includewetlands,montaneandalplne biomes,
Arctic biomes, and coastal biomes. Wetlands will
dry out, wlth grave consequences for amphibians
(Box 7.2), mountainswill become warmer towards
their tops, tundrareglons will warmup, andcoastal
biomes will be flooded. Climaticwarmmg IS
predicted to be greatest at high latitudes. Anlmals
and plants In these reglons will have to cope with
themost rapidchanges. Tundra vegetation may
be pushed northwards as much as 4 degrees of
latitude.This could mean that, for aclimatic
warmlng of 3"C, 37 per cent of present tundra will
become forest. The same degree of climatic warm-
ingwould fuel local temperature increases on
mountains. Animals and plants would need to shift
thelr ranges upwards by about 500 rn. Animal and
plantpopulat~ons on mountains may retreat
upwards as climate warms, but eventually some
of them will have nowhere to go. Thls would prob-
ably happen to the pikas (Othotona spp.) living on
198 COMMUNITY CHANGE

alpinemeadowsonmountainsofthewestern Canadian prairie provinces (Figure 7.13) (Poiani and

United States. Pikas feed on grasses and do not Johnson1991).Itisthemainbreedingarea for
range into boreal forests. Asthe boreal forest habi- waterfowlon theNorthAmericancontinent.The
tat advances up the mountainsides, so the pikas habitat consrstsof relatively shallow-water temporary
will retreat into an ever-decreasing area that may ponds(holdingwater for a few weeksinspring),
one day vanish. This processisalready happen- seasonal ponds (holding water from spring until early
ingtoEdith’scheckerspotbutterfly (Euphydryas summer), seml-permanent ponds(holding water
editha), which lives western NorthAmerica throughoutmuch of thegrowing season inmost
(Parmesan 1996). Records of this species suggest years), andlargepermanentlakes(Plate 7.4). For
that its range is shifting upwards and northwards, breedingpurposes, the waterfowlrequireamixof
and extinctions have occurred at some sites. open water and emergent vegetation. The temporary
and seasonal pondsprovidearich foodsourcein
Communities under pressure the spring and are
used by dabbling ducks. The semi-
permanent ponds provide food and nesting areas for
Community composition and structure change in the
birds as seasonal wetlands dry. As temperatures rise
face of environmentalperturbations.Durmgthe
over the next decades, water depth and the numbers
twenty-first century, many communities will be per-
of seasonal ponds in these habltats should decrease.
turbed by global warming. They will affect the geo-
The lower water levels wouldfavourthegrowth
graphical
location of blomes and
the
species
of emergent vegetation and reduce the amount of
composition of communities.
openwater.Waterfowl may respond by migrating
to differentgeographicallocations,relyingmore
Prairie wetlands in North America uponsemi-permanentwetlandsbutnotbreeding,
Prairiewetlandhabltatoccupiesabroadswath of ot failingtore-nest as theydoatpresentduring
landacross theAmericanMidwestandsouthern droughts.

Pox 7.2
4MPHIBIANS AND GLOBAL frogslivinginwetstream-bankpatches.These
WARMING amphibians retreat into crevices or gather in damp
pockets during the dry season. Four years later, the
4mphibians need water. They must live in it, near golden toad and the harlequin frog had vanished.
t, or else in very humid conditions. Warmer and Rainfall during May 1987 was 64 per cent less than
jrierconditionscould bedisastrousforthem.In average, the lowest ever recorded for that month.
?laces whereglobalwarming is associatedwith Normally, early rainsin May and June helpthe
Increasing aridity, they are likely to suffer. Evidence amphibians to recover fromtheNovember and
af this comes fromendemicgoldentoad (Bnfo December dry season. The 1987 dry season was
kiglenes) and harlequin frog (Ateiopus uavius) popu- extradry.Duringthedrought,watertablesfell,
iationsintheMonteverdeCloud ForestPreserve, springsallbutdriedup,andstreams fedby
Costa Rica(Pounds1990;Poundsand C m p aquifers, which would usually give enough water to
1994). In May and April 1987, there were thou- keep the mossy riverbanks wet, dwindled. As the
sands of golden toads thriving along streams in the amphibiansdisappearedsuddenly,itmay be that
reserve. The maleslivein undergroundburrows theextremedroughtkilledtheadultsandtheir
near thewatertable;theyemergeonlytomate. tadpolesoreggs.Thiscatastropheaugurs illfor
There was also a considerable number of harlequin amphibians during the twenty-first century.
 COMMUNITY
CHANGE 199

Saskatchewan i ,,
,,
Lake ,/'
\ Albefl Vinnrpeg ;
'
I
I ,
\
\
,, Ontario
\
I
I

I
, I

I
I

, I
"
I
I
I
Montana
:I
'.:,*,",." _
\ !"
; Dakota\
North
"""_ ""_ ~
"_
8 """- """"":
I

Y 7 South
Idaho
I
/ i
I
Dakota
I I
/ Wyoming --------__________
I
""
""
-< 1 I "
,".' ........

i
I
I
I Nebraska
1""- I

Figure 7.13 Locaclon of North Amerlcanpralrie wetlands. The prairie wetlands are relatively shallow, water-holding
depressions of glaclal orrgln.
Source: After Poiani and Johnson (1991)

Mires in the Prince Edward Islands

populations at Marion Island, but also prefer larger
TemperaturesinthePrinceEdwardIslands have weevils. A decrease in body size of preferred weevil
Increased by approximately1°C since the early 1950s, prey specles (Bothrumtopusrandi and Ectemnorhinus
while
precipitation has decreased(Chownand similis) hastakenplaceonMarlonIsland (1986-
Smith 1993). The changing water balance has led to 1992), but not on Prince Edward Island. This appears
a reduction in the peat-moisture content of mires to be aresult of increased predation on weevils.
and hrgher growingseason 'warmth'. In consequence, Adults of the prey species, Ectmnorhinus srmilis, are
the
temperature-sensitive and
moisture-sensitive relatively more abundant on Prince Edward Island
sedge, Uncinra compacta, has increased its aerial cover thanadults of thesmallercongener Ectemnorhinus
on Prince Edward Island. Harvestingof seeds by feral m a n o m , and could not be found on Marion Island in
mice (Mus musculus), whichcanstrip areas bare, the late southern summer of 1991. Results not only
has prevented an increase in sedge cover on Marion provlde support for previous hypotheses of the effect
Island. Such extensive use of resources suggests that of globalwarmingonmouse-plant-invertebrate
prey switching may be taking place at Marion Island. interactions on the Prince Edward Islands, but also
Mice not only are eating ectemnorhinine weevils to provide limited evidencefor the first recorded case of
agreaterextentthanfound rn previousstudies of predator-mediated speciation.
200 COMMUNITY
CHANGE

PI- 7.4 Semi- nnanent prairie wetland, Stutsman County, North Dakotu. This wetland is part of
the CoftonwooS(Cake site where long-term hydrological and vegetation data have been collected
b theUnitedStatesGeologicalSurvey(WaterResourcesandBiologicalResourcesDivision).The
prank in the foreground are cattails (Typha spp.)
Photogroph by Karen A. Poiani

Biome and ecotone shifts and 82other.

the
cent
per
in
Temperate
forest
area
alters little. Tropical forests show a slight reduction
Global warming fuelled by a doubling of carbon in area in all but one scenario, which predicts a slight
dioxidelevelscouldproducelargeshiftsin the dis-increase. Maps of predicted change In the leafarea
tribution of biomes. Onestudy,whichexplored five Index (which reflects themaximumrate of trans-
different scenarios forvegetationredistributionwithplration)implydrought-relatedbiomass lossesin
a doubling of atmospheric carbon dioxide levels, pre- most forested regions, even In the tropics. The areas
dicted large spatial shifts, especlally in extratropical most sensitive to drought-induced vegetation decline
regions(Neilson1993a).Largespatialshifis In tem-areeasternNorthAmerlcaandeasternEuropeto
perate and boreal vegetatlon were predicted (Figure western Russia.
7.14). Boreal biomes (talga and tundra) retreat north- In detail, spatial shifts of blomes should produce
wards and decrease in slze by 62 per cent (the range discrete 'change' zones and 'no change' zones (Figure
is 5 1 to 7 1 per cent, depending on the scenario used). 7.15) (Neilson 1993b). Such changes alter the pattern
Boreal and temperate grasslandIncrease In area under of vegetatlon. Large, relatively uniform biomes are
two scenarios. The increase IS 36 per cent one case reduced In m e , while new biomes, which comblne
 COMMUNITY CHANGE 20 1

Area (millions of km2)

-4 0 4 8 12

Non-forest
""""""""

Tropical dry forest

and savannah
"_""""

temperate savannah
"""""""

Temperate forest

Boreal forest
United Kingdom Meteorological Office
UIIl GeophysicalFluidDynamicsLaboratory
0 GeophysicalFluidDynamicsLaboratory,
Q-flux version
El GoddardInstitute of SpaceStudies
Tundra University State N Oregon
a Average

Figwe 7.14 Changes In biome area predicted by the MAPPS (Mapped Atmosphere-PlantSoil System) model under var-
IOUS scenarlos.
Source: After Neilson (1993a)

theoriginalbiomeandtheencroachingbiomes, shouldbecomefragmented as everyvariationin

emerge. Biomes and ecotones alike would probably
affected by droughtfollowed by infestationsand fire if
climate were to change rapidly. Ecotones would be
especiallysensitive toclimaticchange,whether it
wereslow or fast. The pattern of ecotonechange
would be sensitiveto water stress (Figure7.16). With
global warming unaccompanledby water stress, habl-
tats should not fragment as under water stress, but
should display a wave of high habitat variability as
the ecotonegraduallytracks theclimatlcshifts.
Under extreme drought stress, the entire landscape
topography and soil become important to site water-
be
balances and thesurvivorship of different organisms.
The ecotone disappears for a while and reappears at a
new location. It does not visibly shift geographically,
as it would with no water stress, but disassembles and
then re-establishes itself later.
Forest change in eastern North America
It 1s probably common for species to movein the
same general directlon. However, that does not mean
202 COMMUNITY CHANGE

Biome 3

Biome 3

W
1 No
change
Change
Biome 2

p j
-0 zone
2
.- Biome 2 zone
Y

J
Biome 1
........................ ..j......... ... .. ..................:.
; . .. ... ... .
I Biome 1
. .. .. .. .. .. ............:... z
L

.................. *. ....................
!
*: Future
z
\ Present / 'z Transect
Hierarchical level
of change

Fipre 7.15 Geographicalshift in blomes inducedby global warming. Discrete 'change' and 'no change' zones are produced.
Large, uniform biomes shrink. New biomes, which cornbme characterlstics of orlginal biomes and encroachlng biomes,
emerge.
Sorwce: After Neilson (1993b)

thatentirecommunities move together.Quitethe levels. At the year 700,climatestabilizedandthe

contrary, species move at different rates in response
to climatic change. The result is that communities
disassemble, splittmg into their component species
and losing some species that fail to move fast enough
or cannot adapt. Mathematical models are helpful in
understanding the possible changes in communities
as the worldwarms up. Clearly, profoundchanges
in the composition and geography of communities
would occur. This is evidentinthe following case
studies.
An early model simulated forest growth at twenty-
one locations in eastern North America, as far west as
a line joining Arkansas in the south to Baker Lake,
North West Territories,in the north(Solomon 1986).
All forests started growing on a clear plot and grew'
undisturbed for 400 years under a modern climate.
After the year 400, climate was changed to allow for
awarmeratmosphere.Alinearchange of climate
between the years 400 and 500 was assumed, the new
climate at the year 500 corresponding to a doubling
of atmospheric carbon dioxide levels. Climatic change
continued to change linearlyafter the year 500, S O
that, by the year 700, the new climate corresponded
toaquadrupling of atmospheric carbon dioxide
simulation went on for another 300 years. Simulation
runsat each site were repeated tentimesandthe
results
averaged. Validation of the results was
achieved by testing with independent forest composi-
tion data, as provided by pollen deposited over the
last 10,000 years and during the last glacialstage,
16,000 years ago.
Results for some of the sites are set out in Figure
7.17.Atthe tundra-forest border,thevegetation
responds to climatic changein a relatively simple way
(Figure 7.17a). With four times the carbon dioxide
intheatmosphere,theclimatesupportsamuch
increased biomass and,at least at Shefferville in
Quebec, somebirches, balsam poplar, and aspens. The
response of northern boreal forest to warming I S more
complicated (Figure 7.17b).With a four-fold increase
inatmospherlc carbon dioxide levels, theclimate
promotesthe expansion of ashes, birches, northern
oaks, maples, and other deciduoustrees at theexpense
of birches, spruces, firs, balsam poplar, and aspens. In
all northern boreal forest sites, the change In species
composition is similar, but takesplace at different
tlmes. This underscores the time-transgressive nature
of vegetational response to climaticchange.The
 C O M M U N I T YC H A N G E 203

(a) No water stress (b) Water stress

Time 0

'I
I
t

c I Ecotone shifts
I
I
Ecotone disappears

L
Time 2
Ecotone re-established
at a new location
Ecotone shifts again
- .

Distance -
Figure 7.16 Ecotone changes induced by global warmlng. (a) With no water stress. (b) With water stress.
S o r o w : After Neilson (1 993b)

southern boreal forest and northern deciduous forest
(Figures 7 . 1 7 ~andd) also have complicated
a 0. Biomass declineseverywhere,generally
response toclimatlcchange.Inbothcommunities,
species die back twice, the first time around and just
after 500 years, and the second t m e from about 600
to 650 years. In the southernboreal forest, the change
is from a forest dominated by conifers (spruces, firs,
and pines) to a forest dominated initially by maples
and basswoods, and later by northern oaks and hicko-
ries. At some sites in the northern deciduous forest,
species appear as climate starts to change thenvanish
once the stable climate associated with a quadrupled
level of atmospheric carbon dioxide becomes estab-
lished. Specles thatdothls Include thebutternut
Vzqhluns c-znereu),black walnut (Jugluns nzgru), eastern
hemlock (Tsrtgu cunrrdensrs), and several species o f
northernoak.
In western and eastern deciduous
forests, the response of trees to climatic warming I S
remarkably uniform between sites (Figure 7.17e and
as soon
as warming starts in the year 400. The drier sites in
the west suffer the greatest losses of biomass, as well
as a loss of species; this is to be expected as prairie
vegetation would takeover. The declineof biomass in
the eastern deciduous forest probably results chiefly
from the increased moisture stress in soils associated
withthe warmer climate.The increased moisture
stress also gives a competitive edge to smaller and
slower-growing species, such as the
southern
chlnklpin oak (Quereus muebhlenbergii),post oak (Quereus
stellutu), and live oak (Quercus virginzutu), the black
gum (Nyssu sylvutrca), sugarberry (Celtrs lumigata),
and the American holly (Ihlex opacu), whlch come to
domlnatetherapid-growlng species such as the
American chestnut (Custuneu dentutu) and various
northern oak species.
204 COMMUNITY C H A N G E

1 Present-day
climate
l$l :+, 1
N
4 x ~
climate

( b ) Northern boreal forest

~ 2

120

80

40

160
(c) Southernborealforest (d) Northern deciduous forest
I I I I

1b o
I ( e ) Western deciduous forest

80

40

0
500 500 1,000
Time (years)
 C O M M U NCI T
HYA N G E 205

Soil water regimes and forest change
Forest growth is sensitive to changesinsoilwater
regimes.Globalwarming is suretoalter water
regimes and this will have an inevltable impact upon
forests. A forest-growth model was used in conlunc-
tion with a soil model to assess the impact on forests
of a doubling of atmospherlc carbon dioxide levels
(Pastor and Post 1988).The modelwas run for several
sites in the north-eastern United States, including a
site in north-eastern Minnesota. The climate of this
region is predicted to become warmer and drier. The
major changes will occurat the boundarybetween the
boreal forest and cool temperate forest. Forest growth
was modelledon two soil types:soils withahigh
water-holding capacity and soils with a low water-
holding capacity. Thesimulations beganin 1751.
Bare plots were 'sown' with seeds of the tree species
common in the area. Themodel forest was then
allowed to grow for 200 years under present climatlc
conditlons. Next, carbon dioxide concentratlons were
gradually increased until, after century,
a their
present value had been doubled. After having reached
that value, carbon dioxide levels were held constant
for the next 200 years.
Themodelpredictedthat on soils withahlgh
water-holding capacity, where there will be enough
water available to promote tree growth, productivlty
and biomass will increase. O n soils with a low water-
holding capacity, productivity and biomasswill
decrease in response todrierconditions.Inturn,
raised productivity and biomass will up levels of soil
nitrogen, whereas lowered productivity and blomass
will down levels of soil nitrogen.Thevegetation
changes are,therefore,self-relnforcing. O n water-
retentive soils, global warming favours the expansion
of northern hardwoods (maples, birches, basswoods)
at the expense of conifers (spruces and firs); on well-
drained, sandysoils, it favours the expansion of a
stunted oak-pine forest, a relatively unproductive
vegetatlon low in nitrogen, at the expense of spruce
and fir.

Figure 7.17 Simulatlons of forestbiomassdynamlcs over one millennium in response to climatic change induced by
lncreaslng levels of carbon dioxlde In the atmosphere at SIX sltes In eastern North Amerlca. (a) Shefferville, Quebec (57"N,
67"W). (b) Kapuskaslng, Ontario(49"N, 83"W).(c) West upper Michigan (47"N, 88"W). (d) North central Wisconsin
(45"N. 90'W). (e) South central Arkansas (34"N. 93"W). (0 Central Tennessee (36"N. 85"W). The tree specles are as
follows: A. American beech (Fagrrs grundi/&a); R. American chestnut (Castanea denfafa);C . American holly (Ilex opacu); D.
ashes: green ash (Fraxrnrrs penxrylc,un/ru), whlte ash (Fraxrnrrsutr,ertcatru), black ash (Fruxrtrusnrgra), blue ash (Fraxrnlrs
yrrudratrgrrlaru);E. basswoods:Amerlcanbasswood (Tilia atuerrcanu) and whlte basswood (Tilia beteropbylla); I:. blrches:
sweet birch (Betula letrtu), paper birch (Betula papyri/ira), yellow blrch (Bernla ullegbatrrensrs), and gray blrch (Betula pop-
trlijinlia); G . balsam poplar (Pop~lus ba/sut~~$eru). bigleaf aspen (Po/url/rs grundidetrtata),trembling aspen (Poprr1rr.rtremdozdes);
1 3 . black cherry (Prrrw.r .rero/nru);I . black g u m (Nyssa s y / r ~ / ~ c aJ.
) ; butternut (Jtrghns crtrereu) and black walnut ( J q l a n s
nmgru);K . eastern hemlock (Tsfrga catrudetrrrs); L. elms: Amerlcan elm (U/?uwaruerrcanu) and wlnged elm (Ultmrs alutu); M.
firs: balsam fir (Ab2e.r holsrrt~rea)and Fraser fir (Abiesfraseri);N . hlckories: bltternut hickory (Caryu c-ordiforttm).mockernut
hickory (Carya tomntosa), plgnuc hlckory ( C a v a glabra), shagbark hickory ( C a v a n t ~ / r r ) shellbark
, hickory (Carya lucru-
rosrr), andblackhickory (Carya t e x a m ) ; O. hornbeams:easternhornbeam (Ostrya zwgrmana) andAmericanhornbeam
(Curprnrr.r cwo/inrn?ra);P. maples: sugar maple (Arer sacrburrrtn),red maple (Acer rubra),and silver maple ( A msaccharmum);
Q. northern oaks: white oak (Qtrerurs all~u),scarlet oak (Qnerctrs c.occmnerr), chestnut oak (Qtrercusprrnus), northern red oak
(Qrrrrc~rs rtdra), black oak (Quercx.r rrlrr/ma), bur oak (Qrrercrrs macrocurpa), gray oak (Qtren.rr.r borealis), and northern pln oak
(Querc~rsellipso/dulis); R. northern whlte cedar (Tbtrp occtdenta/i.r), redcedar (Jlrnzpernsvrrgmzunu), and tamarack (Larmx
larmtru); S. plnes: jack pme (Pitrzrs bnnkszrrna),red plne (Pinm resrwoso), shortleaf plne (Pintrs ec.brna/a),loblolly plne (Pinrrs
iaedu), Virginla pine (Pinus t~rrgm~una), and p t c h plne (Pinus rmgtdu), and, 'I', white plne (Piwr.r s/roBtrs);IJ. yellow buck-
eye (Aewrlrrs ortundra); v spruces: black spruce (Pil-eu tmrrana), and red spruce (Picea r/rbens);and v l whlre spruce (Piceu
glarrca); w. southern oaks: southern red oak (Qtrercxsfalcatrr), overcup oak (Qnere-nslyruta), blacklack oak (Qtrercmrsmarr-
landia), chinkipln oak (QIMWS tmreblenbergii),Nuttall's oak (Qrrerc-usnuttallii), pin oak (Querempuhstrts), Shumard's red
oak (Qtrwcxr shrrmardii,post oak (Qrwwr srellata), and live oak(Qmmws rwgmnzann);X. sugarberry (C'eltls lurt,rga/a);Y.sweet-
g u m (Lzqtrrdumbur sryracrfltra);Z. yellow poplar (Lrr/odendrot/ tdip$eru).
Source: After Solomon (1986)
206 COMMUNITY CHANGE
Disturbance and forest dynamics
Global warming would favour a rise in the rate of
forest disturbance owing to an increase in meteor-
ological conditions likely to cause forest fires
(drought, wind, and natural ignition sources),con-
vective windsandthunderstorms, coastalflooding,
and hurricanes. Simulationssuggestthatchanges
in forest composition associated with global warming
woulddependuponthedisturbanceregime(Over-
peck et a/. 1990).Twosets of simulations were
run.Inthe first set (‘step-function’experlments),
simulated forest was grown from bare ground under
present-day climate for 800 years. Thisenabled
the natural variability of the simulated forest to be
characterized. At year 800, a single climatlc variable
was changed In a slngle step to a new mean value,
which perturbed the forest. The simulation was then
contmued for a further 400 years. In each perturba-
tionexperiment,theprobability of acatastrophlc
disturbance was changed from 0.00 to 0.01 at year
800. This is a realisticfrequency of aboutone
plot-destroying fire every 115 years when a 20-year
regeneration period(during whlch no further cata-
strophe takes place) of the trees in a plot is assumed.
In each of the step-function simulations, three types
of climaticchange(perturbation) were modelled:
a 1OC increase in temperature; a 2°C increase in tem-
perature; and a 15 per cent decrease in precipitation.
Inthe second set of simulationruns(‘transient’
experiments), forest growth was, as in thestep-
function experiments, started from bare ground and
allowed to run for 800 years under present climatic
conditions.Then, from the years 800 to900,the
mean climate,bothtemperatureandprecipitatlon,
was changed linearly year by year tosmulatea
twofold increase in the level of atmospheric carbon
dioxide,untilthe year 1600 whenmean climate
was again held constant. As In thestep-function
experlments,theprobability of forest disturbance
was changed from 0.00 to 0.01 at year 800. In all
slmulationruns, a relatively drought-resistant soil
was assumed,andthe resultswereaveragedfrom
40 random plots into a single time series for each
model run.
Themodel was calibrated forselected sites in
the mixed coniferous-hardwood forest of Wisconsin
and the southern boreal forest of Quebec. Selected
summary resultsare presented in Figure7.18.An
increase in forest disturbance will probably create a
climatically induced vegetation change that is equal
to, or greaterthan,thesameclimaticallyinduced
change of vegetation without forest disturbance. In
many cases, thisenhancedchangeresulting from
increased disturbance is created by rapid rises in the
abundances of species associated with the early stages
of forest successlon,which appear because of the
Increased frequency of forest disturbance.Insome
cases, as in Figures 7.17a, d, and e, a step-function
change of climate by itself does not promote a signifi-
cant change in forest blomass, but the same change
workinghand-in-handwith Increased forest distur-
bance does have athoroughgoing effect on forest
compositionand biomass. Interestingly, the altered
regimes of forest disturbance, as well as causinga
change in the composition of the forests, also boost
the rate at which forests respond to climatic change.
For instance, in the transient climate change experi-
ments, where forest disturbance IS absent through the
entire duration of the simulation period, vegetation
change lags behind climatic change by about 50 to
100 years, andsimulatedvegetationtakesat least
200 to 250 years to attain a new equilibria1 state.
In thesimulationruns where forest disturbance
occurredfrom year 800 onward,thevegetatlonal
change stays hard on the heels of climatic change and
takes less than 180 years to reach a new equilibrium
composition after the climatic perturbation at year
800.
Ecological lessons from simulation models
Species show a wide range of responses to climate.
In consequence, the response of different animal
and plant species to climatic change will be varied.
Thlsshould mean thatnaturalcommunities are
likely to disassemble and that habitats restructure in
transient,
a non-equilibrium fashion as climatlc
change unfolds. Validated models that help forecast
theseeventsare needed to aid scientists in better
 COMMUNITY CHANGE 207

160

80
m
f
5 0
c
.-m0
L
1601 I With increase
disturbance I 1
80

0
a White pine $"1Oaks Early successional
trees
- 0
Hickories
Maples trees Other

80
m
f
5 0
I With disturbance
increase I 1

0 0 00 0 00 0 0
0 0 0 0 0 0
co N co N co
Time (years)
Jack pine Firs White pine
0Birches Spruces n
hardwoods
Other

F i p 7.18
~ Simulated changes in species composition of forests at two sites investlgated in eastern North America. (a)
to (c) is a site in Wisconsin, and (d) to (0 is a site in southern Quebec. At both sites, experlments were run with an
increase in disturbance at year 800 (top of figure for each slte) and without an increase In disturbance at year 800 (bot-
tom figure for each site). Additlonally, three climatic change scenarios were slmulated: l0Catemperature Increase at year
800 (left-hand figures, a and d); a 15 per cent decrease In preclpltation (mlddle figures, b and e); and a translent change
in which mean monthly preclpttatlon and temperature were changed linearly from year 800 to year 900 and thereafter
held constant (nght-hand figures, c and f).
Souwe: After Ovetpeck et al. (1990)
208 COMMUNITYC H A N G E
understandingthe e c o l o g d ramifications of global E S S A Y Q U E S T I O N S
climatic change.Also, and perhaps more Important for
conservationbiology,suchvalidatedmodels can help 1 Why hasClements‘sunidirectional view
provide probabilities for the occurrence of these
events, which will allowpolicy makers to make better,
2 Why does habitat fragmentation pose a
informeddeclsions (T. L. RootandSchnelder 1093).
SUMMARY
Communities change. The nature of this change I S
debatable. Theclasslc view saw climatic climaxes and
balanced ecosystems resulting from unldirectional
successlon. Succession 1s a complex process and IS
explained by at least three models - the facilitatlon
model,the tolerance model, and
the
inhibltion
model. It may also be drlven by fiactors external to the
community (allogenic factors). Prlmary succession is
the colonization of land or submarine surfaces that
have never existed before. Secondary successlon is the
Invasion of newly created surfaces resulting from
removal of pre-existmg vegetation. A modern view
of communitychange stresses thedisequilibrium
behavlour of communitles. It sees succession going
inmanypossible directions,and sees communities
as temporary collections of specles that assemble
and disassemble as the envlronment changes. Much
communltychange has been caused by land cover
transformation over the past two hundred years. Two
Important aspects of this transformation are habitat
fragmentation,and I t attendant effects onwildlife,
andthe loss of wetlands.Globalwarmingduring
the twenty-first century is likely to put peripheral,
geographicallyrestrlcted,highly specialized, poorly
disperslve, andclimatically sensitivespecies under
intense pressure. It will also cause significant changes
in many communitles. Wetlands, tundra, and alpine
meadows are especially vulnerable.
Of succession been revised?
threat to many species?
3 What communitychanes are likely to
occur as a result of gloI 2al warming?
F U R T H E RR E A D I N G
Committee on Characterization of Wetlands,
National Research Council (1995) Wetlands:
Charactertstics and Boundaries, Washington,
DC:
National Academy Press.
Gates, D. M. (1993) Climate Change and Its Biological
Conseqwences, Sunderland, Massachusetts:
Sinauer
Associates.
Huggett, R . J. (1993) Nodefling the Httnzan Impact on
Nutwe: Systenu Analysis of Envtronnmtal Problems,
Oxford: Oxford University Press.
Huggett, R. J. (1997) EnvironmentalChange: The
EvofLhg Ecosphere, London: Routledge.
Jackson, A. R.W.andJackson,J.M. (1996)
Environnrental Scrence: The Natltral Envtronment and
Ht/n~anImpact, Harlow: Longman.
Matthews, J. A. (1992) The Ecology of Rerently-
Dqlaciated Terrain: A Geoecologtcal Approarb t o Glacw
Forelandr and Primary Succession, Cambridge:
Cambridge Universlty Press.
Peters, R. L. and Lovejoy, T. E. (eds) (1992) Global
Warnmg and Bzological Dtversity, New Haven,
Connecticut and London: Yale Unlversity Press.
Schwarrz, M. (1997) Consewation In Highly Fragmented
Landrtapes, New York: Chapman & Hall.
 LIFE, H U M A N S ,A N DM O R A L I T Y
Human attitudes toulards the living world are relevant to biogeographers involved with
conservation a n d ecosystem management. This chapter covers:
non-humanrights
kinds of environmentalism
biogeographicaltheoryandconservationpractice
Biogeographical and ecological prlnciplesnaturally
Inform questions of envlronmental management.
However, envlronmental management does not just
have a ‘sclentific’ dimenslon. It has social and ethical
dimensions, too. All dimensions must be consldered
in discussing the relations between humans and other
livlngthings.Thischapter will explore therlghts
of animals and Nature, attitudes towards Nature, as
seen in the different brands of envlronmentalism, and
therelationships between biogeography, ecology,
and conservation practlce.
BlORlGHTS
Environmental ethics deals wlth value judgements
aboutthe biologlcal and physicalworlds. I t was
begun, at least in Its modern form, by philosophers
during the 1970s. Environmental ethics is a diverse
andimmense field of study. I t arose from three
conslderatlons (Botkin and Keller 1995: 619). First,
human actlons, aided by technologlcal developments,
affect Nature deeply, so it seems necessary to exam-
ine the ethical consequences of these actions. Second,
many human actions have world-wde consequences,
and a global perspective ralses Its own moral ques-
tions. Third, moralconcerns have expanded so that
animals, and even trees and landscapes, possess moral
and legal rights - civilization includes all Nature
wlthln its ethicalsystems. This expansion of concerns
demands a new environmental ethics.
Animal wrongs: do organisms have
rights?
What are rights?
Rights are a social contrivance that help people to
live together. They are not some quality that indivrd-
uals possess. Rather, they are a quality that members
of a socletyare willing to pretend that individuals
possess. Such apretence assures socially acceptable
behaviour (most of the time). This idea is straight-
forward, though open to philosophical debate. The
big questlon is whether the scope of ‘rights’ shouldbe
extendedtoincludeother species, and even rocks,
rlvers, and landscapes. There I S no convincmg reason
why they should not be so extended. If rlghts should
help humans to avoid conflict with one another, why
should not animal rights, plant rlghts, and landscape
rights help humans to protect thelr envlronment?
An all-encompasslngdefinitlon of rlghts would
helphumansto respect Nature.Butthematter is
complicated by three preconditions, without whlch
210 LIFE, HUMANS, AND MORALITY
an objectcannot be grantedrights - commensur-
ability, responsibility, and sensibility (Tudge 1997).
Commensurability means that rightsare a reciprocal
agreement - you respect my rights and I’ll respect
yours. Unfortunately, this equitable arrangement is
distorted by power structures. Some groups of people
have less power to infringe the rights of others. More
powerful groups (the richer for instance),therefore,
tend to possess more rights than less powerful groups
(the poorer for instance). As animals, plants, and land-
scapes have no effective power, why should they be
grantedrights?Stated so bluntly,thlsargument
smacks of insensitlvity. To sootheconsclences, a rider
is added - to earn rights, people must accept respon-
sibility.
Animals,plants,
and landscapes cannot
accept responsibility, therefore they can have no
rights. However, there IS no reason why rights and
responsibilitles should be Inseparable. If animals are
worthy of human respect, then they can be awarded
rights, even though they should have nofeelings
toward humans.Sensibility, or lack of It, inall
species save humans, arose from a mechanlstic v ~ e wof
animal behaviour instigated by Ivan Pavlov and hls
dogs. To be brief,most scientists are now prepared
to see otheranlmals as somethmgmorethanblo-
chemicalmachlnes. I t wouldseem that ‘respectable
science has caught up wlth theanlmal-lovers’ and I t IS
no longer perverse ‘to perceive animals as sentient
beings, who register their emotional responses with
varylngdegrees of consciousness’ (Tudge 1997: 41;
see also Regan 1983).
So, if animals have rights, how should they be
treated? This is a difficult Issue with many aspects.
All the aspects stem from a sea change in human
attitudes toward animals, at the root of which is a
new-found sympathy towards the natural world, and
towards anlmals in particular (e.g. Fisher 1987). This
sympathy has drlven the forces of animal welfare over
recent years. Theanimalliberationmovement,
through films andothermedia, has shownpeople
how many anlmals are forced to live in pounds, on
factory farms, and In laboratories.People have been
stirred into various klnds of actlon - boycotting pet
shops, breakinginto laboratories to free animals,
ceaslng to eat meat, spurnlng the wildlife trade, and
so on.However,thechange in humanattitudes
toward animals does not express itself solely in
militant action. There is a qulet slde to the change
that is less evident but possibly more fundamental.
Two examples from AustraliaandNew Zealand
(O’Brlen 1990) should demonstrate this subtle and
yet profound attitude change.
In Australia, the VictorianAcclirnatlzation Society
was formed in 1861. A central objective of this early
wildlife management society was ‘the introductlon,
acclimatizarlonanddomesticatlon of allinnoxious
animals, birds, fishes, Insects and vegetables whether
useful or ornamental’(Rolls1969).This objective
was regarded as worthy and servlng the public inter-
est by sclentists, natural historians, and landholders
of the Soclety. InmodernAustralia,theacclimati-
zation of exotic specles would be regarded by many
as Irresponsible and bizarre, and is prevented by
legislatlon. Attitudes towards ’pests’, some of which
are the ancestors of the ‘innoxious’ animalsintro-
duced in the nineteenth century, have also shifted.
Take the example of the rabbit. The huge Australian
rabbltpopulation was controlled in the1950s by
introduclng myxomatosls. But In 1987, when the
same vlrus was consldered as a control agent for the
wild rabbit population I n New Zealand, the welfare
of wild rabblts was put first and the virus was not
introduced.
Three moral issues arising from thenotion of
animal rlghts deserve closer scrutiny - culling
animals, the wildlife trade, and keepmg animals in
captivity.
Culling
Controllinganimalpopulatlons may mean culling.
Few people raise objectionstothe eradication of
viruses, bacteria, or pestilential insects. Attempts to
limit populations of organisms higher up the evolu-
tionary ladder cause eyebrows to be raised. Mammal
culling causes public outcry. The cullingof seal pups
IS met by a vociferous and appalled response.
Theshooting of feral horses from helicopters
began In Australia in late1985. By 1988,the
CommonwealthDepartment of PrimaryIndustries
 LIFE, H U M A N SA, N D
MORALITY 211

was receiving more than 7,000 protest letters a year, and to ban trade in a further 800 specles threatened
mostly ‘campaign’ mail from correspondents overseas with extinction.
(O’Brien 1990). In the United States, feral horses are About a quarter of the wildlife trade is unlawful
protected by law. The Wild Free-Roaming Horse and commerce in rare andendangered species. The un-
Burro Act (197 1)decrees that feral horses and burros fortunate animals are poached in the wild and smug-
must be consldered as an integral part of the natural gled across frontiers. A typlcal year will see the sale
systems in which they are found. The Actprovides for of 50,000 primates, tusk ivory from 70,000 African
the Bureau of Land Management to set suitable levels elephants, 4 million live birds,10millionreptile
for herd size, and for surplus animals to be removed skins, 15 million pelts, 350 million troplcal fish, and
and offered for privatemamtenance by qualified about 1 million orchids (Lean and Hinrichsen 1992:
individuals, or destroyed if they are old, sick, lame,or 145).Peoplecaught illegallytrafficking In wildlife
private care cannot be found. In 1987, 8,000 horses are arrested, fined, and even jailed. Thegrowing
were held in captivity awaitlng ‘adoption’. willingnessanddeterminationtostopthewildlife
Many people accept thatsomehumaneculling trade is anotherinstance of humans upholding the
of largemammals is a necessary evil toprevent rights of anmals.
over-exploitation of vegetatlonandenvtronmental
degradation. In
GreatBritain, for example,the
Keeping animals in captivity
culling of deer populations is probably necessary.
With animal rights now to the fore, the method of Zoologicalgardens are notwhat theywere. As
culling I S the subject of scientific study. On Exmoor society’s attltudes toward animals have changed, zoos
and the Quantock Hills, In Somerset, red deer (CKWLS have responded by rethinkingthe reasons for their
K ~ U ~ ~ are
U J culled
) by rifle (stalking) and by hunting existence. Zoos began in the heyday of natural
with hounds. A study carried out for the National history, when almost every educatedVictorian was
Trust(the Bateson Report)concludedthathunts an amateur naturalhlstorlan.Interestsatthetime
with hounds cause deer great distress. The exerttons centred around collectmg, identifying, and naming.
of the chase change muscles and blood far beyond Zoos were places to display the ordinary and bizarre
anychanges that would be expected In normal life. creatures found in far-flung parts of the globe. This
Red deeraresedentary animals that escape natural role for zoos lingered well into the twentieth century.
predators (mainly the wolf, although there are none Even in 1970, the lion enclosure at Newquay Zoo, in
now in Britaln) by brlef sprints. Hunting deer with Cornwall,England, was designedto keeplionson
hounds is not ‘natural’, because wolves would never display to the public for as long as possible. This was
chase deer for longdistances. As a result of the achieved by keeping them in a relatively small space,
Bateson Report, membersof the National Trustvoted and letting them return to their even smaller sleep-
to ban deer hunting with hounds on all its lands. ing quarters only at nlght. In the 1990s at the zoo a
largelionenclosureincorporatesmanyfeatures that
resemble the lions’ natural savannah environment.
The wildlife trade
There is, for instance, a platform (simulating a small
The wildlife trade is an international business knoll) and trees. Food is hidden around the enclosure
worth about $15 billion a year. Much of it I S legal and the lions have to find It. And the lions are free to
and controlled by national laws and an international ‘go indoors’ when they wlsh.
treaty - CITES (Convention on International Trade A brochure for Newquay Zoo captures the splrlt
inEndangered Specles of Wild Fauna and Flora). of the tlmes when i t says that good zoos are good
CITES was drawn up in 1975 and 139 countries are for anlmals and good for people. They are good for
now party to it. Collectlvely,these countrles act to anmals for one reason only - conservation. Five
regulate and monltor trade In around 25,000 spec~es aspects of conservation are paramount:
212 LIFE, H U M A N S , A N D M O R A L I T Y

Breeding
1 programmes save animals from Does all Nature have rights?
extinctlon.
2 Wildlife research helps to preserve habitatsand
A belated effect of the Darwinian revolution is that
save animals. humansare now seen as part of Nature.Environ-
3 Animal behaviour research assists their survival in mental philosophers now passionately prosecute this
the wild. idea, but remain unclear as to its implications. What
4 Providingsanctuaries for animalsthat have lost does it mean to be part of Nature? This philosophi-
their habitat. cal issue is complexand beset by aterminological
5 Provlding hospitals for sick and injured animals. labyrinth (see Colwell 1987). But all commentators
agree that ‘being part of Nature‘ dispenses with the
Unlesswild populations are supplemented by duality between humans and the natural world, pro-
captive breeding, and culling populations that out- vldes a moral justification for treating Nature more
humanely, and gwes a philosophical ~ustification for
grow their reserves, then most of the world’s largest
terrestrlal mammals will vanish. Zoos make signifi- seeing intrinsic valueinall things(e.g.Callicott
cant contributions to conservatlon as genetic refuges 1985; McDaniel 1986; Zimmerman 1988).
and reservoirs, especially for large vertebrate species An early and eloquent advocate of theintrinsic
worth of the natural world was Aldo Leopold (1049),
threatened with extinction (Rabb 1994).Przewalski’s
who advanced an all-embracing land ethic. Leopold
horses ( E ~ u przeutafskii)
J are believed to be extinct
In thewild;thecurrentknownpopulation of 797 affirmed the right of all resources, including plants,
animals existsin zoos (Boyd 1991).It is hoped to animals, and earth materials, to continued existence,
remtroducethis species toits former Mongolian and, at least In places, tocontinued existencein a
habitat by the early years of the twenty-first century. natural state. This land ethic assumes that humans
Zoo resources for conservation and research are are ethically responsibility tootherhumansand
limited, so zoos are encouraglngthedevelopment human societies, and to the wider envlronment that
includes, animals, plants, landscapes, seascapes, and
of criteria to help prioritize actions for conservatlon
the air. Granting rlghts of survlval to animals does
of biodiversity.NorthAmerican,European,and
Australian zoos are assisting the developmentof tech- not necessarily mean that they cannot be eaten, but
it does mean that endangered spectes should be cared
nical capacltles among zoo counterparts, government
agencies, andprotected areas in bothdevelopmg for and helped to recuperate. A land ethic behoves
and developed countries of the world to further the humans to sustain Nature for the present generation
conservation of biodiversity. and for future generatlons.
The question of Nature’s rights arose as a legal
A remarkablesuccess story (so far) I S the goldenlion
tamarin ( L ~ O W Z ~ ~roJalia)
Z ~ J (p.197).Thlssquirrel- issue in the 1970s. Mineral King Valley is a wilder-
sized primate lives in themuch-reduced lowland ness area in the Sierra Nevada, California.Disney
Atlantlc forest, Brazil. Since the mid-l99Os, a com- Enterprises, Inc., wlshed to develop the valley as a
binatlon of relocating five tamarin familiesfrom skiresort withln multimilliondollar recreational
vulnerable areas Intotheprotectlon of the POGO facilities. The Sierra Club (founded in 1892 to help
das Antas Biological Reserve, and relntroduclng preserve theYosemite Valley and other
natural
147 captive-bred tamarins into the wild, has put the wonders In California) objected that the development
little prlmate on the road to recovery. For long-term would destroytheaesthetlc value and ecologlcal
survival, the tamarm’s habitat area will need to be balance of the area, and brought a suit against the
government. The Sierra Club could not claim direct
doubled.
harm from thedevelopment,andthe land was
government owned and the government represented
the people, so peopleingeneral were not wronged.
 LIFE, H U M A N S , A N D M O R A L I T Y 213

A lawyer, ChristopherD.Stone (1972), suggested brands tend to polarize around, at one extreme, eco-
that, as there were precedents for Inanimate objects centrism, with Its decidedly motherly attitude
such as shlps havlng legal standing, trees should also towards the planet; and, at the other extreme, tech-
have legal standing. So the sult was made on behalf nocentrism with a more luissez-farre, exploitative
of thenon-human wilderness. The case was taken attitude (Table8.1). The ‘green’ endof this spectrum
tothe US SupremeCourtbut was turneddown. was Initiated by theatomlcbombandgalvanued
However, in a famous dissenting statement, Justice into action by pesticide abuse. Each strand represents
William 0. Douglas proposed the establishment of a a system of beliefs, although no individual would
new federal rulethatwould allow ‘environmental necessarily believe in just one strand.
issues t o be litlgated before federal agencies or federal
courts in thename of theInanimateobjectabout
Technocentrics (pale greens and very pale
to be despoiled, defaced, or invaded by roads and
greens)
bulldozers and where injury is the subject of public
outrage’. Although trees were not given legal rights Technocentrics tend to have a human-centred vlew
in thls case, theethlcal values and legal rights for of the Earth coupled with a managerial approach to
wilderness were discussed. resource development and environmental protection.
They favour maintaining the status quo in existing
governmentstructures.Technocentricstendto be
A T T I T U D E ST O W A R D S N A T U R E conservatlve politlclans of all political partles, leaders
of industry, commerce andtrades’ unlons, and skilled
The Age of Ecology opened on 16 J d y 1945 In the workers. In short, they are members of the produc-
New Mexican desert ‘wlth a dazzling fireball of light tive classes. They aspire to improve wealth for them-
and a swelling mushroom cloud of radioactive gases’ selves and for society at large. They en~oymaterlal
(Worster 1994: 342). Observmg the scene, a phrase acquisition for own sake and for the status I t endows.
from the Bbagazlarl-Gita came Intoproject leader And they are politically and economlcally very
J. Robert Oppenhiemer’s mind: ‘I am become Death, powerful.
the shatterer of worlds’. For the first time in human There are two brandsof technocentrlc - the cornu-
history, a weapon of truly awesome power existed, a copians (or optimists) and the accommodationists
weapon capable of destroying life on a planetary scale. (or environmental managers).Conservative techno-
From under the chillingblack cloud of the atomic age centrics are cornucopians or optmlsts.They are
arose a new moral concern - envlronmentalism - that named after the legendary goat’s horn thatoverflowed
soughttotemperthemodern science-basedpower with fruit, flowers, and corn, and signified prosperity.
over Nature with ecological inslghts into the radia- They have faith in the applicatlon of sclence, market
tion threat to the planet. The publication of Rachel forces, and managerial ingenuitytosustalngrowth
Carson’s Silenr S p r r q (1962), the first study to bring and survlval.
the lnsldious effects of DDT applicationtopublic Liberal technocentrics are accommodationists.
notice, was more grist for theenvlronmentalist’s They have falth in the adaptability of institutions and
mill. The dual threats of radiation and pesticldes set mechanisms of assessment and decision makingto
environmentalism moving. accommodate envlronmental demands. They believe
that adjustments by those In positions of power and
significance can meet theenvlronmentalchallenge.
Brands of environmentalism
Theadjustments involve adaptlngtoandmould-
There are several brands of environmentalism, each ing regulation (including environmental tmpxt
of whlch promotes a different attitude towards the assessment) and modifylng managerial and busmess
envlronment (see O’Rlordan 1988, 1996). These practices to reduce resource wastage and economlcally
2 14 LIFE, H U M A N SA,N M
D ORALITY

Table 8. I Brands of environmentalism and their characteristics

Environmental Type and colour Characteristics

inclination

Ecocentrism Deep ecologists or Stress the intrinsic importance of Nature for humanity
(Naturecentred) Gaians (dark greens) Believe that ecological (and other) laws should dictate
human morality
Ardently promote biorights - the right of endangered
species, communities, and landscapes to remain
unmolested

Communalists or Emphasizethesmallness of scale ('small is beautiful') and

self-reliance hence
community
identity in settlement, work, and leisure
soh-technologists Interpret concepts of workand leisure through a process
(medium greens) of personal and communal improvement
Stress the importance of participation in community
affairs, and guarantee of the rights of minority interests

Technocentrism Accommodationists Believe thateconomic growthand resource exploitation

(humancentred)or
environmental
can
continue assuming: suitable economic adjustments to
managers(pale taxes,
fees,
etc.;
improvements in the legal rights to a
greens) minimum level of environmental
quality;
compensation
arrangements satisfactory to those who experience
adverse environmental and social effects
Accept new project appraisal techniques and decision
reviews arrangements to allow for wider discussion or
genuine search for consensus among representative
groups of interested parties

Cornucopiansor Believe that humans canalwaysfind a way out of any

optimists (very pale difficulty, beit political, scientific, ortechnological
greens - dark blueAccept that prc-growth goalsdefine the rationalityof
under the surface) project appraisalandpolicy formulation
Optimistic about the human ability to improve the lot of
the world's people
Faith that scientific and technological expertise provides a
firm foundation for advice on matters pertaining to
economic growth, public health, and public safety
Suspicious of attempts to widen basis for participation
and lengthy discussion in project appraisal and policy
review
Believe that all impediments can be overcome given a
will, ingenuity, and sufficient resources arising out of
growth

inconvenientpollutlon,without any fundamental
shift in the distribution of political power. Pressure
groups aligned to technocentrics tend to be NIMBY
(not-in-my-back-yard)groups - theydonotmlnd
development so long as ~t does nottake place near
them - and private Interest groups (consumer groups,
civil liberties groups, and so on). They are very vocal
and command much media attentlon.
Ecocentrics (medium greens and dark greens)
Ecocentrics have a Nature-centred vlew of the Earth
in which social relations cannot be divorced from
people-mvironmentrelations.They hold a radical
vlslon of how future soclety should be organized. They
would give far more real power to communlties andto
confederations of regionalInterests. Central control
and natlonal hegemony, so treasured by techno-
centrlcs, are the very antithesls of ecocentrism. Eco-
centrics tend to be found among those on the frlnge
of modern economles, that is, the‘non-productlve’
classes - clerics, artists, teachers, students, and, to an
ever increasing extent, women. Theyare characterized
by willingness
a to eschew materlal possessions
for their own sake, and to concern themselves more
with relationships; andby a lack of faith in large-scale
modern technology and assoclated demands on elitlst
expertise and central state authority. Pressure groups
aligned to ecocentrics tend to be green groups and
alternativesubcultures(e.g.
femlnists,
the peace
movement, religious movements).
There are twobrands of ecocentrics. The more
conservative ecocentrlcsare communalists or self-
reliance, soft technologists. They believe in the co-
operativecapabilities of societies to be collectlvely
self-reliant using approprlate science and technology.
That I S , they believe In theability of peopleto
organlze theirown economles if glventherlght
incentlves and freedoms. Thls vlew extends a ‘bottom
LIP’ approach to the development of the Third World
based on the application of indigenous customs and
appropriate technical and economicassistance from
western donors.
Liberal ecocentrics are Gaians or deep ecologists
(Box 8.1). They believe in the rights of Nature and
LIFE, HUMANS, A N D MORALITY 215
the essential co-evolution of humans and natural phe-
nomena. Extreme ecocentrics believe that the moral
basis for economic development must lie in the inter-
connections between natural and social rights: there
is no purely anthropocentric ethic. Within thls group
might be Included a powerfulreligiouslobby (Box
8.2).
Alignments of ecologists
I t would be wrong, if excusable, to Imagine that all
ecologists andenvironmentalscientists hold eco-
centric views. Therelatlonship betweenecocentric
environmentalists and ecologists has not always
been as cosy as might be supposed: ecologists do not
always say what ecocentrlcs wantto hear. Paul
Colinvaux (1980: 105) wrote: ‘If the planners really
get hold of us so that they can stamp out all indi-
vidual liberty and do what they like wlth our land,
they might declde that whole countles full of Inferior
farms should be put back into forest’. His displeasure
wlth land-use planningandenvironmentalism IS
evident. Later In the book (Colinvaux 1980: 1lo), hls
words smack of social Darwinism and ‘Nature red
in tooth and claw’, when he talks of different specles
‘going about earnlng thelr livlngs as best they may,
each In its own indivldual manner’, and what ‘look
likecommunity properties’ being ‘thesummed
results of all these blts of private enterprise’, though
elsewhere he sees peaceful coexistence, not struggle,
as the outcome of natural selection, the peace belng
broken In upon by a devlant aggressor species such as
Hotno .rappiens who seek to encroach on another’s niche
(Colinvaux 1980: 131). Today, few ecologists whole-
heartedlyacceptColinvaux’s viewpolnt,and many
approach Naturegreen In head andheart.One of
the many I S Danlel B. Botkin (1990) who advocated
uslngmodern technology In a constructwe .and
positive manner, a position that tries to brldge the
mlddleground betweenecocentrlsm andtechno-
centrlsm.

- ~~~~
Table 8.3 Different attitudes of deep and shallow ecologists
Environmental
Shallow
ecology
problem
Pollution Advocate a technology that seeks to
purify the air and water and to spread
pollution more evenly; laws limit
possible pollution; polluting industries
are preferably exported to developing
countries
Resources Emphasize resources for human
beings, especially for the present
generation in affluent societies.
Resources belong to those who have
the technology to exploit them.
Resources will not be depleted
because, as they get rarer, higher
market prices conserve them, and
substitutes will be found through
technological progress. Plants, animals,
and natural objects are valuable only
as resources for humans; if they should
have no use to humans, they can be
destroyed
Population See overpopulation as a problem of
developing countries. The question of
the optimum human population is
discussed without reference to the
optimum population of other life forms.
The destruction of wild habitats is
accepted as a necessary evil
Cultural diversity Regard industrialization of the kind
and appropriate manifested in the west as an example
technology for developing nations
Source: After Naess (1 986)
LIFE, HUMANS,ANDMORALITY 2 17
Deep ecology
Evaluate pollution from an
ecospheric point of view, not
focusing on its effects on human
health, but on the effects on life as 1
a whole, including life conditions
of every species and ecosystem
Deeplyconcernedwith resources ~
and habitats for all life-forms for
their own sake; no natural object
is regarded solely as a resource;
this view leads to a critical
examination of human modes of
production and consumption
Recognize that excessive pressures
on planetary life stem from the
human population explosion. The
pressure stemming from industrial
societies is a major factor, and
population reduction in these
societies, as well as in the
developing countries, is
imperative
Concerned with maintaining
cultural diversity, and limiting the
impact of western society upon
presently existing non-industrial
societies, defending the fourth
world against foreign domination,
and applying local, soft
technologies where appropriate
2 18 LIFE, HUMANS, AND MORALITY

Box 8.2
RELIGION, NATURE, AND THE command, builds an ark to save species onEarth
WORLD WILDLIFE FUND from extinction when God sends a grand Flood to
punish humanity. After the Flood has finished, God
Inautumn 1986, auniquealliance wasforged makes a covenant - t h e first in the Bible. He makes
between conservation, as represented by the World hiscovenantnot just withNoah,butwithall
Wildlife Fund, and five of the world’s chief reli- creatures on Earth, in the seas, and in the skies. God
gions. The venue was Assisi, in central Italy, home promises never wantonly to destroy life again, and
of thepatronsaint of animals.Apilgrimage, sets the rainbow in the sky to bear witness to his
conference, cultural festival, retreat, and interfaith promise.
ceremony in the Basilicz of St Francis formed the TheRainbowCovenant has beeninspired by
basis of a permanentbondbetweenconservation God’s covenantwithNoah,and was specifically
and religion. designed by the InternationalConsultancy on
Forreligions,ecologicalproblemsposemajor Religion, Education and Culture (ICOREC) to be
challengesinrelatingtheologyandbelieftothe usablebyanyone,religious or not. So, unlike
damage to Nature and human suffering caused by the Hittite and biblical covenants, it is unilateral -
environmental degradation. All too ofien, religious a contrite pledge by humans to other living beings
teaching hasbeenused or abusedtojustifythe and their ecosystems. The rainbow’s arc embraces
destruction of Natureandnatural resources. In not only those who live, but those who will live; it
the declarations issued at Assisi, leading religious encompasses all living creatures and the earth, air.
figuresmappedoutthe way to re-examine and and water that sustain them. This is the Rainbow
reverse this process. Andin 1987, the Baha’i’ Covenant:
Movement, which had its origins in Islam, joined Brothers and Sisters in Creation, we cwenant this aky wid
the World Wildlife Fund’s New Alliance. At the you and with all mation yet to be;
same time, starting with the conference and retreat With awy living mature and all that contains andsustain.
heldin Assisi,religiousphilosophersarehelping you.
With allthat is on Earth and within the Earth itself;.
inject
somepowerful
moral
perspectives
into
Withall thatlives in thewaters and with the water
conservation’s ill-defined ethical foundations. themselves;
Adevelopment of the Assisieventwas the With all that pies inthe skies and with the sky itseq
making of the Rainbow Covenant at Winchester We establish this covenant, that all our PWS will be US^
Cathedral,EnglandinOctober 1987. Covenants to prevent your destntction.
We confus that it is our own kind who p:tt yon at risk 4
have long
a history. For theancientHittites,
dpath.
covenants were made between rulers and subjects for We ask fw your t m t and as a symbol of our intention i ~ ’
the benefit of both. For Jews and Christians, ‘The mark this cmwant with you by the rainbow.
Covenant’ is between God and his people, and was This is the sign of the covenant betum ourselm and
made after the Flood. In the Bible, Noah, at God’s living thing that is found on the Earth.

A balancing act: biorights, human Ecosystem management arose in the late 1980s
rights, and ecosystem management and is advocated by many scientists and other people
Do humans still have rights in the world of the deep interested In the environment (e.g. Agee and Johnson
ecologist? People are partof Nature. The philosophy 1988). Its ultimate aim is to enhance and to ensure
behind a new brand of conservation - ecosystem the diverslty of species,communities,ecosystems,
management - accepts that simple fact. and landscapes. It is a fresh and emergmg model of

resource management. It covers spectrum
a of
approaches. Amild,technocentric version simply
extends multiple use, sustained yieldpolicies,pro-
secutingstewardship
a approach, in which the
ecosystem I S seen merely as ahuman life-support
system (e.g. Kessler et al. 1992). In this v~ew, public
demands for habitat protection, recreatlon, and
wildlife uses are simply seen as constraints to maxi-
mizing resource output (Cortner andMoote 1994). A
more radical, ecocentric approach is to accept Nature
on itsownterms, even where doing so means
controlling ~ncompatible human uses (e.g. Keiter and
Boyce 1991). This extreme, but eminently sensible,
form of ecosystem management reflects a willingness
to place environmental values, suchas blodiversity and
animal rights, and social and cultural values, such as
the upholding of human rights, on an equal footing.
It means accepting that environmental concerns are
not purelyfactual; they have do withvalues and ethics.
Peopleexert a profound influence on ecological
patterns and processes, and in turn ecological patterns
and processes affect people. The connection between
technology and the environment is well studied; not
so the connectionbetween social system and the envi-
ronment. However,policies are tending to move away
from the administrator-as-neutral-expert approach
to policies that engender public deliberation and the
discovery of shared values. Naturally, such extension
of ecolog~cal matters to the soc~al and political arena
presents difficulties, though these may not be in-
superable (e.g. Irland 1994). Ecosystem management
accepts that human values must play a leading role in
policy decision-making. Conservation strategies must
takeaccount of human needs andaspirations;and
they must integrate ecosystem, economic, and social
needs. The key players in ecosystem management are
scientists, policy makers, managers, and the public.
The public, many of whom have a keen interest ~n
environmental matters, are becoming more involved
in ecosystem management as professlonals recognize
the legitimacy of claims that various groups make
on natural resources. In Jervis Bay, Australia,the
marine ecosystem is used by many existingand
proposed conflicting interests (national park, tourlsm,
urbanization,militarytraining)(Wardand Jacoby
LIFE, H U M A N S , A N D M O R A L I T Y 219
1992). Similarly, in the forests of the south-western
United States, ecosystem, economic, and social needs
are considered in policy decision-making concerning
ecology-based, multiple-use forest management
(Kaufmann et a / . 1994) (Figure8.1).
B I O G E O G R A P H Y ,E C O L O G Y ,A N D
C O N S E R V A T I O NP R A C T I C E
Biogeographical and ecological ideas inform conser-
vation practice. When ruling theories ~n ecology and
biogeography change, conservation
practice
soon
follows sult.This
generalization,
though
valid,
shouldbetreatedcautiously, as therelationships
between ecological science and environmental policy
are complicated (Shrader-Frechette and McCoy
1994). Nevertheless, it is fair to say that the major
paradigmshifts in twentieth-century ecological
thought have engendered quite distinct conservation
philosophles. Four paradigms, which were mentioned
in Chapter 7, are recognized: endurlng equilibrium,
balancedecosystems, disequilibriumcommunities,
and the edge of chaos.
Environmental exploitation and
enduring equilibrium
Environmental problems in the offing
In his Walden;or, Liji in the Woods (1893 edn), Henry
Davld Thoreau expressed concern for wild Nature and
wild environments. H e urged that, when a conflict
arises between Nature and human society, then the
first thing to consider are the laws of Nature. George
Perkins Marsh wrote ManandNature;or,Physical
GeographyasModified by Human Action in 1864. H e
examined‘thegreater,morepermanent,andmore
comprehensive mutations which man has produced,
and I S producing in earth, sea, and sky; sometimes,
indeed,with conscious purpose,but for themost
part, as unforeseen thoughnatural consequences of
actsperformed for narrower and more immediate
ends’ (Marsh 1965 edn: 19). He believed that ‘Man is
everywhere a disturb~ng agent. Wherever he plants
220 LIFE, HUMANS, AND MORALITY

Social
needs
I

ecosystem and human

benefits, costs, and risks Economic and social needs
(National Environmental
Policy
Act) + + + + ,
I
’
I

Ecological principles
Ecological principlesfilter
I
Biological
Species integrity
loss unrest
Ecological,
economic,
Implementation
Economic
socialDegraded
and
environments
sustainability
instability
4
Monitoring

and feedback

Figure 8.1 T h e Integration of ecologlcal, economtc, and social needs In a decwon-analysis model. Economlc and social
needs are tested agalnst an ‘ecologlcal filter’, whlchIS shown In the shaded box. The aim IS to determine economlc and
social actlons that will produce the most deslrable balance between blologlcal Integrity and ecological, economic, and
soclal sustainability Bowlng fully to economlc and soclal needs would lead to specles loss and envlronmental degrada-
tion. Bowlng fullyto ecological needs would leadto soclal unrest and economlc Instability A compromlse posltlon allows
the malntenance of blological Integrity while catermg for economic and soclal needs. The resulting decision model leads
to the implementation of an envlronmental policy.The effects ofthe policy are carefully monitored and evaluated. Ifthe
policy should fail to work as desired, then amendments can be made and the process started anew, until a satlsfactory
outcome IS achieved.
Source: After Kaufmann et al. (1994)
 LIFE, H U M A N S , A N D M O R A L I T Y

hisfoot, theharmonies of natureareturnedto land use. Thispractlce, heaverred, shouldstop

discords’(Marsh 1965 edn: 36). Marsh was the first environmental degradation andplace the entire nation
modern scholar to see humans as a factor, and not on a biologically and economically sustainable foot-
merely as a subject, in Nature. ing.Inother words, hewas proposing ecological
In1922,Robert Lionel Sherlock published an princlples to gulde conservation practlce and achleve
enormously important book entitled Man as a a relationship with Nature that ensured an enduring
Geological Agent: A n Aa-otmt o f His Artton o n lnanitnute equilibrium.
Nattrre. From extensive observatmns withtn
the
British Isles, Sherlock showed the effects that human
Balanced ecosystems
activities were havingon the landscape, the effects
that forestry, grazlng, agricultural management, and The balanced ecosystem view, and its extension in the
Gaia hypothesis, supplanted ‘enduring equilibrium’
the technologlcal and d u s t r i a l activity of twentleth- as
century society were havingonlandscape processes. a blueprintfor Nature’s design.I t guided conservation
Specifically, he looked at the effects of road and rail- strategy in two different ways. First, ecocentric envi-
way construction, open-cast coal minlng, and slate, ronmentalistsarguedthatunrestratned interference
stone, gravel, and sand quarrying; and the manage- withNature’sdevelopmentstrategy leads to eco-
ment of waterways and coasts. He gathered starlstlcs system damage,andthatthe world’s endangered
glving the amounts of material extracted or removed, ecosystems should be defended against the excesses of
and furnished estimates of the rates of erosion and human actlons. ‘Spaceship Earth’, a term first used
sedimentatlon. by Adlai Stevenson, a Unlted StatesAmbassador, in a
During the late 1920s and the 19.30s. the recogni- speech given before the United NatlonsEconomic and
tion of the human Impact on the soil and the need for Social Council in Geneva on 9 July 1965, became a
soilconservatlon was a pressingconcern,largely leading environmental idea.
because of the dust-bowl that devastated the Great Theimage of Spaceship Earth has now lost Its
Plains reglon In the Unlted States. The main problems
potency. Nonetheless, i t sparked off an envlron-
of environmental degradation were found m a d y in mentally frlendly brand of economics. Free enterprlse
western Europe and North Amerlca. and Marxist economists placed humans at the stage
centre and tended to disregard the natural environ-
ment, seelng it as a free and inexhaustible reservoir
Broken equilibrium
and a bottomless rubbish dump (Cottrell 1978: 7).
Whatever theroot cause ofenvlronmental degradation This exploitative attitudefostered traditional resource
should be, Clements’s notton ofenduring equilibrium management practice that maximizes productmn
suggested that humans had erred. Its message for con- of goods and services through sustained yield from
servation was clear: the climax state is Nature’s deslgn balancedecosystems; adoptsutilitarian values that
for bioticcommunities so humansshould respect regard human consumptlon as the best use of
and preserve it. I t was thought that the equilibrium resources; and holds a continuous supply of goods for
betweenhumansandNature had been upset and human markets as the purpose of resource manage-
should be restored, that Nature’s deslgn princlples had ment(CortnerandMoote 1994). Such barefaced
been forsaken and needed Immediate relnstatement. ‘resourclsm’ IS ecologically unsound. I t fails to recog-
Thls response toenvlronmentaldegradatlon was nize limltstoexplomtionand, in consequence, a
evldent in Paul Sears’s Daert.r on the Alurch (1949, the growingnumber of species, and even entire eco-
first edition of whlch appeared In 19.35, In the after- systems, are currently endangered. But, unsound or
math of the Amerlcan dust-bowlcalamity. Sears not, I t persists. Even now, ecosystems are vlewed by
advocated that each county In theUnitedStates some as long-lived, multi-product factories (Gottfrled
should appoint an ecologist to adv~se on matrers of l992), o r , if you prefer, as Nature’s superstores.
222 LIFE, H U M A N S , A N D M O R A L I T Y
the
In
mid-l960s,a new ‘environmental
economics’ emerged that was sensitive to the needs
of the environment. Kenneth Ewart Boulding (1966,
1970), for instance,complainedthathumans have
operated a ‘cowboy economy’,in which success is
measured in the amount of material turned over and
in which resources are exploited in a profligate and
reckless manner. Instead, he insisted, life on a small
and crowded planet demanded a‘spaceship economy’,
inwhich success is gauged by turnoverandthe
watchwords areconservation, maintenance,and re-
cycling.Eugene P. Odumemployedthe spaceship
analogy as recently as 1989. He recalled the crisis on
board the spacecraft Apollo 13. An explosion in the
craft had caused damage to the vessel and threatened
the lives of the crew.After several anxious hours,
the crew managedto reach the safety of thelunar
module and abandoned ship. Odum compared this
event to the current environmental crisis on Earth.
Our life-support systemsarebeingdestroyed,but,
unlike the Apollo 13’s crew, humans cannot abandon
ship and find a safe haven. The only option is to stay
onboardSpaceshlp Earthand repair thedamage.
Unfortunately, he said, humans do not know how to
repair the planet, although he hinted that a civiliza-
tion capable of putting people on the Moon should
have witenoughtocomprehendthe mechanlcs of
ecospheric dynamics and keep the cogs of the eco-
logical machine running freely. Luckily, he observed,
the Earth has its own regeneration systems, its own
means of healing wounds. The view that the Earth
has itsown powers of healing is supported by the
Gaiahypothesis,the medicalanalogy beingmade
clear in Lovelock’s (1989) use of the term ‘geophysi-
ology’. (As an aside, I wouldpointoutthatthe
impossibility of indefinitely sustaining life in a
spacecraft was exploredin Brian Aldiss’sfictional
‘Helliconia’ trilogy, which also used an exotic setting
tospeculate on thepossibilities of the Gaia hypo-
thesis.)
Second, technocentric environmentalists used the
balanced ecosystem view of Naturetopromote
the understanding of balanced ecosystem function as
a basis for planetary management. Both ecocentrics
andtechnocentricsthoughtthey could determme
what was safe for humans to do and what was not.
Largely, they were mistaken (see Worster1994:
416).
Many ideas of theequilibrium ecologlstsare
used today. Equilibrium ecological thought also
underpinsthe theory of islandbiogeography, as
formulated by Robert H. MacArthurandEdward
0. Wilson in the 1960s, which is still used to aid the
designing of nature reserves. Some conservationists
have espoused the steady-state Gaia hypothesis as a
blueprint for planetarymanagement (Myers 1987),
and somevery green environmentalists have used it as
a design for living. The equilibria1 ecological theme
persists in the notion of sustainability. This is the
well-meaning, but possibly misguided, idea that the
ecosphere should be developed without compromis-
ing the ability of future generations to satisfy their
needs. Or, put another way, sustainable development
urges that diverse, functioning ecosystems should
be preserved withoutdamagingthe economy; and
that economies and human welfare need preserving
as much as theintegrity of ecosystems (Gerlach
and Bengston 1992). It was first presented in 1980 at
an internationalforum in theWorld Conservatlon
Strategy, by the Internatlonal
Unlon for the
Conservation of Nature(IUCN),and has become
mainstay of much environmentalist philosophy. The
IUCN identified three key areas for action: the main-
tenance of essential ecological processes and life-
support systems, thepreservation of genetic diversity,
andthesustainable use of species and ecosystems
(Allen 1980).
But the idea of sustainability is questionable. One
problem lies in integratinghumanand biophysical
factors. Humans interact
with
Naturethrough
culture, often in symbolic ways that are not compre-
hended by biological or physical ecosystemmodels
(Gerlach and Bengston1992). In other
words,
humans can generate wants and capabilities of meet-
ing these wants that lie outside the naturalecological
order. This is often difficult for scientists to under-
stand, and leads to their focusing on envlronmental
protectionandrestoration,ratherthan facing the
greater challengeof understanding social and cultural
interactions with thebiophysical world. Thevery Idea

of sustainability itself is a curious human construct.
Laudable though the idea of sustainability might be,
there are problems with it, not the least of which is
the lack of a definition.It was originally taken to
mean meetingthe needs of the present generation
without compromising the ability of future genera-
tionsto satisfy their needs (World Commission on
EnvironmentandDevelopment1987),butat least
eight interpretatlons are available (Gale and Cordray
1991).It is therefore a buzzword that lacks bite.
Anotherproblem is its
inadvertent arrogance -
shouldthe present generation be so presumptuous
and foolhardy as to anticipate the needs of future
generatlons?Thesepithyquestionsshouldnot be
ducked In ecosystem management initiatives
Evolutionary disequilibrium
Some time In the 197Os, the notions of homeostasls,
balance, and stability i n Nature collapsed. They were
supplanted by the idea of evolutionary disequilib-
rium. Thedramatlcchange of paradigms was, In
part, a reflectlon of the human soclety at that time. I t
arose when i t wasGashionable to see neither Nature
nor society as a stable entity,steadfastly wlthstanding
all attempts to dislodge it from a global equilibrlum
state. Instead,a view emerged of all hlstory as a record
of disturbancespringingfromboth‘cultural and
naturalagents,
including droughts,
earthquakes,
pests, viruses, corporate takeovers, loss of markets,
new technologies, increaslng crime, new federal laws,
and even the invasion of Amerlca by French literary
theory’ (Worster 1994: 424, emphasisin original).
Disequilibrium ecologists seemed divided among
themselves on the advice they should give to society
about how to act over the environment. One group,
reflecting some of the new disequilibrium thlnking,
began to challengethe
publicperception
that
ecology and environmentalism were the same thing.
Some ecologlsts became disenchanted with trying to
conserve a healthy planet. Nature is characterized by
highlylndivdualistic assoclatlons, so why attempt
to constrain I t ? This anarchlc argument, if taken to
theextreme, could have revlved social Darwinism
andstood in antitheslstothe conservation ethic of
L I F E , HUMANS, AND M O R A L I T Y 223
co-operation and collective action suggested by
Odum’s balanced ecosystems. In the event, a mildly
anarchic view of Nature was taken by some ecologists
(p.215).Anothergroup of ecologists, instark
contrast,drew differentconcluslons from thedis-
equilibriumtrendswithinthendiscipline.Daniel
B. Botkin(1990) was one of themostarticulate
advocates of a new, chastened set of environmentalist
policies. H e favoured an environmentalism that was
more friendly towards manipulating and dominating
Nature,butthat tolerated modern technology and
progress, and human deslres for greater wealth and
power. This was not the preservation of a balanced
nature, so much as a responsible attitude to techno-
logical developments and their effects on an environ-
ment that would inescapably change. Botkin’s specific
proposals were somewhat vague. He cautioned that we
should not engineer Nature at an unnatural rate nor
In novel ways. But In a worldwhere disturbance is
commonplace and change is the rule, how may the
unnaturally rapld andthe novel be identified and
defined!
The edge of chaos
The implications of chaotic ecosystem dynamics
for conservation are far from clear. How are conser-
vationists to use a chaotic design? Answers to this
question have to confront a paradox: a chaotic view
of Nature I S at once exhilaratingandthreatening.
ChaoticNature, so irregularandindividualisticin
character, appears almost impossibly difficult to
admlre or to respect, to understand or to predict. It
seems to be a world In which the security of stable,
permanent rules IS gone forever, a dangerous and un-
certain world that Inspires no confidence (Prigogine
andStengers1984:2 12-1 3). Thisdark aspect of
chaos might promote a feeling of alienation from the
naturalworld,and cause peopletowithdrawinto
doubt and self-absorptlon (cf. Worster 1994: 413). It
mlght also setpeople wondering how theyshould
behave In a world where chaos reigns. With natural
disturbance foundeverywhere,why shouldhumans
worry about doing their own bit of disturbing? Why
not join individuals of all other specles and do their
224 LIFE, H U M A N S , A N D M O R A L I T Y
own thlng, free of guilt that in doing so they would
cause any specific damage? As Worster (1994: 413)
put it, what does the phrase 'envlronmental damage'
mean In a world so full of natural upheaval and
unpredictability?
However, chaos does not have to be portrayed in
such gloomy and doom-laden terms. Chaotlc Nature
has abrlghtandedifylngaspect,too.In a chaotic
world,communltles, ecosystems, and socletiesare
sensitivetodisturbance. Small disturbances can,
sometimes,growand cause the communities, eco-
systems, or societies to change. Consequently, i t is a
world In which indivldualactlvlty may have major
slgnificance (cf. Prlgogine and Stengers 1984: 3 13).
It is apost-modern worldinwhlch Increased Incli-
vidualityand diversityencourages agreat overall
harmony (a notlon akin to Taoist beliefs). Moreover,
the newfangled theory of chaotic dynamlcs is leading
to the discovery of hiddenregularities In natural
processes, the application of whlch is prowng most
salutary (Stewart 1995). Satellites can now be sent to
new destlnations, ferried by gravitational forces
through a fractal Solar System, with far less fuel than
was once thought possible. A dishwasher, involvlng
tworotatingarmsthatspln chaotically, has been
invented in Japanthat removes dirt fromdishes
using less energy than in a conventlonal dishwasher.
Some forests and fisheries are being better managed
through an understanding of thelr chaotic behaviour.
And, chaos theory is glvlng us a breathtaking view of
our Universe, and persuading us to look again at our
place wlthin It. It offers us a new design with which
to reconsider our world, and a new framework wlthln
which to review our fears and our hopes.
Forward from the past: the
biodiversity bandwagon
By the last decade of twentieth century, ecologlsts
were undecided about
the
implications of their
subject for modern technological civilization. None-
theless,theyfoundthemselves regrouplng around a
new conservation ideal - biodiversity. I t mattered
notthatNature was chaotlcandunpredictable.It
was also gloriously diverse, and, for all its disturbing
and perverse ways, and its abiding ability to evade
our understanding, I t still needed our love, our
respect, and our help (cf. Worster 1994: 420). This
V I ~ Whas supplied a conservation ethlc, and a source
of inspiration (and funding) for biogeographers,
for the closing years of the twentieth century, and
perhaps beyond.
SUMMARY
There aremoral grounds for grantlngrightsto
animals, plants, andeven landscapes. In the late 1990s
attltudes towards anlmals are more sympathetic than
they were In the past. Thls sympathy I S seen in the
publicoutcryagalnst cruel methods of culling, in
the measures taken to stop the wildlife trade, and in
the new role for zoos In conservation. Attitudes
towards Nature are vaned and reflected in different
brands of environmentalism. The two main opposing
brands ofenvironmentalismare ecocentrism and tech-
nocentrism. Ecosystem managementaccommodates
human needs andaspirationswlthin anecocentric
perspective.Biogeographical and ecological theory
mforms conservation practice. During the twentieth
century the leadingideas in biogeography andecology
have changed.In consequence,conservation guide-
lines have changed. As the twenty-first century
draws near, the chaotic world of the theoretical ecol-
o p s t stands parallel with the blodiverse world of the
practlcal ecologist.
E S S A YQ U E S T I O N S
Should animals, plants, and landscapes
be granted rights?
To what extent does biogeographical
and ecological theory inform
conservation practice?
What is so special about ecosystem
management?
 LIFE, H U M A N A
SN, MDO R A L I T Y 225

FURTHER READING

Anderson, E. N . (1996) Ecologies of the Heart: Emotfon.

Belid and the Enzironnrent, New York: Oxford
Universlty Press.

Botkm, D. B. (1990) Discordant Harmonies: A Neur

Ecology for the Tuznty-First
Century, New York:
Oxford University Press.

Commlttee onScientific Issues in the Endangered

Species Act, National Research Council (1995) Scienle
and the Endangered Spec.ies Al.t, WashingtonDC:
National Academy Press.

Cooper, N . andCarling,R.C. J. (eds) (1996)

Ecologists and EthicalJudgernents, London: Chapman &
Hall.

Norton, B. G. (1991) Touwd Untty Among

Environmentalists, New York:
Oxford University
Press.

Ryder,R. D. (ed.) (1993) Anutral Welfare and the

Envirotment, London: Duckworth and RSPCA.

Shafer, C. L. (1990) Nature Reserves: Island Theory and

Consetvatton Practtte, Washlngton and London:
Smlthsonlan Institunon Press.

Sylvan, R.andBennett,D. (1994) The Greening of

Ethm: From Hnmun Charwnurn t o Deep-Green Theory,
Cambridge: Whlte Horse Press; Tucson, Universlty
of Arlzona Press.
 GLOSSARY

Table GI The geological time-scale for the last 570 million years

Era and subera Period Epoch Age range (millions of year ago)

Start Finish

Cenozoic era
Quaternary sub-era Pleistogene Holocene 0.0 1 0
Pleistocene 1.64 0.01
Tertiary subera Neogene Pliocene 5.2 1.64
Miocene 23.3 5.2
Palaeogene Oligocene 35.4 23.3
Eocene 56.5 35.4
Palaeocene 65 56.5
Mesozoic era
Cretaceous Late 97 65
Early 145.6 97
Jurassic Late 157 145.6
Middle 178 157
Early 208 178
Triassic Late 235 208
Middle 24 1 235
Early 245 24 1
Polaeozoic era
Permian 290 245
Carboniferous 362.5 290
Devonian 408.5 362.5
Silurian 439 408.5
0rdovician 510 439
Cambrian 570 510
 GLOSSARY 227

abiotic Characterized by the absence of life; in- atonal soils Soils in which erosion anddeposition
animate. dominate soil processes, as insoilsformedin river
alluvium and sand dunes.
acidophiles Organisms that love acidic conditions.
bacteria Micro-organisms, usually single-celled, that
adaptation The adjustment or the process of adjust-
exist as free-living decomposers or parasites.
ment by which characteristm of an entire organism,
or some of its structures and functions,become better barrier Any terrainthathinders or preventsthe
suited for life in a particular envlronment. dispersal of organisms.
aerobicDependingon, or characterized
by, the basal metabolic rate The rate of energy consump-
presence of oxygen. tion by an organism at rest.
algae Simple, unicellular or filamentous plants. behaviourThe reactlon of organismstoglven
circumstances.
alkaliphilesOrganismsthat flourish in alkaline
environments. benthic Pertalning to the bottom of a water body.
allelopathicPertainingtothe influence of plants biocideA poison or othersubstance used tokill
upon each otherthroughtoxicproducts of meta- pests (and inadvertently other organisms).
bolism - a sort of phytochemical warfare.
bioclimatesTheclimaticconditionsthat affect
allogenic Originating from outslde a community or living things.
ecosystem.
biodiversity The diversity of specles, genetic infor-
anaerobicDependingon, or characterizedby, the mation, and habltats.
absence of oxygen.
biogeochemical cycles The cycling of a mineral or
anthrax An infectious and often fatal disease caused organic chemical constituent through the biosphere;
by the bacterium B a c i l h antbraczs; common in cattle for example, the carbon cycle.
and sheep.
biologicalcontrolThe use of natural ecological
apical buds Buds located at the tips of shoots. lnterrelatlonships to control pest organisms.

aquifer Asubsurface geological formation contamng biomagnification

Theconcentration of certain
water. substances up a food chain.

archipelagoAgroup of islands,typically a large biomantle The upper portion of soil that is worked
group. by organlsms.

aridity The stateor degree of dryness.
atmosphereThe gaseousenvelope of theEarth,
retained by the Earth’s gravitational field.
autogenic Originating from inside a community or
ecosystem.
autotrophsOrganisms - greenplantsandsome
mlcro-organisms - capable of making their own food
from inorganic materials.
available moisture Precipitation less evaporation.
biomass The mass of living materlal In a specified
group of animals, or plants, or in a community, or in
a unit area; usually expressed as a dry weight.
biome A community of animals and plants occupy-
ing a climatically uniformarea on a continental scale.
biosphere Life andlife-supporting systems - all
livingbeings,atmosphere,hydrosphere,and pedo-
sphere.
biota All the animals(fauna) and plants (flora) living
in an area or region.
228 GLOSSARY

biotic Pertalning to life; anlmate. cohort Indivlduals In a population born durlng the
same perlod of t m e , e.g. during a partlcular year.
biotic potential The maximum rate of population
growth that would occur In the absence of environ- commensalism An interaction between two species
mental constraints. In which one specles (the commensal)benefits and the
other specles (the host) is unharmed.
borealforest A plantformation-type associated
wlthcold-temperateclimates (cool summersand communityassemblyThecomingtogether of
long wlnters); also called talga and coniferous ever- species to form a communlty.
green forest. Spruces, firs, larches, and pines are the
competitionAn interactlonbetweentwospecies
domlnant plants.
trying to share the same resource.
browsers Organlsms that feed mainly on leaves and
competitive exclusion principle The rule that two
young shoots, especially those of shrubs and trees.
species wlth Identical ecological recluIrements cannot
bryophytesSimple landplantswlthstemsand coexist at the same place.

leaves but no true roots o r vascular tissue: mosses, congener An organism belonging to the same genus
hornworts. and liverworts.
as another or others.
cactiNewWorldplantsbelonging to the family conifers Trees of the Order Coniferales, commonly
Cactaceae with thick, fleshy. and often prickly stems. evergreen and bearing cones.
calcicoles (calciphiles) Plants that love soils rich in consumption Community respiration.
calcium.
continentaldriftThe differential
movement of
calcifuges (calciphobes) Plants that hate soils rlch contlnents caused by plate tectonlc processes.
in calcium.
corolla The inner envelope of a flower; I t IS made of
capillary pressure The force of water in a capillary fused or separate petals.
tube.
corridors Dispersal routes offerlng little or no rem-
carrion Dead and decaying flesh tance to migratlng organisms.

carrying capacity The maxlmum population that an Cretaceousperiod A slice of geological tlme
environmentcan
support
without
environmental spanning 145.6 million to 65 million years ago; the
degradation occurrlng. youngest unit of the Mesozoic era.

caviomorph rodents A suborder of the Rodentla cullingThe act of removing o r killing‘surplus’

Cenozoic era A slice of geological time spannlng 65
million years ago to the present; the youngest unlt of
the geological eras.
chasmophytes Plants that live in crevlces.
chomophytes Plants that live on ledges.
chronosequence A time sequence of vegetatlon or
soils constructed by uslng sltes of different age.
animals In a herd or flock.
cursorial Adapted for running.
cuticle A protecttve layer of cutin (a wax-like, water-
repellent materlal) covering the epldermls (outermost
layer of cells) of plants.
cycad Any gymnosperm of the Family Cycadaceae
looklng like a palm tree but topped with compound,
fern-like leaves.

circulatory system The system ofvessels through decomposer An organism that helps to break down
which blood is pumped by the heart. organic matter.
 GLOSSARY 229

demeA group of interbreeding Indivduals; a local El Niiio Theappearance of warm water in the usually
population. cold water regions off the coasts of Peru, Ecuador,
and Chile.
density-dependent factors Causes of fecundity and
mortality that become more effectlve as population endangered species A specles faclng extlnctlon.
density rises.
endemicspeciesA species nativeto a particular
density-independentfactors Causes of fecundity place.
and mortality that act independently of populatlon
environmentThe biological (biotic)and physical
density; natural disasters are an example.
(ablotic)
surroundings
that Influence Individuals,
detritivoreAnorganismthatcomminutesdead populations, and communities.
organic matter.
environmentalethicsA philosophy dealingwith
detritus Dislntegrated matter. theethics of theenvironment,includinganimal
rlghts.
diatom A microscopic,unrcellular, marine(plank-
tonic) alga with a skeleton composed of hydrous environmental factor Any of the biotic or abiotic
opaline silica. components in the environment, such as heat,
moisture, and nutrient levels.
dicotyledonous plant species Plants belonglng to
the Dlcotyledoneae, one of the two major divisions environmental gradient A continuous change in an
of flowering plants. They are characterlzed by a pair environmental factor from one place to another; for
of embryonic seed leaves that appear at germination. example, a change from dry to wet conditlons.
disharmonious community community
A of environmentalism A movementthatstrivesto
animals and plants adapted to a climate that has no protect the environment against human depredations.
modern counterpart.
Eocene epoch A slice of geologlcal time spanning
dispersal The spread of organisms into new areas. 56.5 million to 35.4 million years ago.
dispersalrouteThepath followed by dispersing epiphyte A plant that grows on another plant or an
organlsms. object, uslng I t for support but not nourishment.
distributionThegeographical area occupied by a estuarine Of, pertaining to, or found In an estuary.
group of organlsms (species, genus, family, etc.); the
same as the geographical range. evaporation The diffusion of water vapour Into the
atmosphere from sources of water exposed to the alr
d r o u g h t A prolonged period with no rain.
(cf. evapotranspiration).
ecological equivalents Species of different ancestry
evapotranspiration Evaporation plus
the water
but with the same characteristics livlng In different
dischargedintotheatmosphere by plant
trans-
biogeographical regions. Also called vicars.
piration.
ecosphere The global ecosystem - all life plus its
life-support
systems (air,
water,
and soil). evergreen A plant
with foliage that stays green all
year round.
ecosystem Short for ecolog~calsystem - a group of
organismstogetherwlththe physical environmentextinctionThedemise of a specles o r any other
taxon.
with
interact.
which they
ecotone A transirlonal zone between twoplant extirpation The local extinction of a species or any
communities. other taxon.
230 GLOSSARY

extremophiles Organisms that relish ultra-extreme and comprising large parts of South America, Africa,
conditions. India, Antarctica, and Australia.
fauna All the animals living in an area or region. grazer An organism that feeds on growing grasses
and herbs.
fecundityReproductivepotential as measured by
thenumber of eggs,sperms, or asexual structures gross primary production
Production before
produced. respiration losses are accounted.
filter route A path followed by dispersing organisms habitat The place where an organism lives.
that,owingtotheenvironmental or geographical habitatselectionThe selection of particular
a
conditions, does not permlt all species to pass. habitat by a dispersing individual.
flora All the plants living in an area or region. heath A tract of openand
uncultlvated land
food chain A simple expresston of feeding relatlon- supporting shrubby plants, and especially the heaths
shipsinacommunity,startingwithplantsand (Erzcu spp).
ending with top carnivores. heliotropism The growthof a plant towards or away
food web A network of feeding relationships within from sunlight.
a community. helophytes Marsh plants.
frugivorous Pertaining to fruit-eating. heterogeneous An environment comprising
a
mosaic of dissimilarspatialelements;non-uniform
GaiahypothesisThe idea thatthe chemical and.
environment.
physical conditions of the surface of the Earth, the
atmosphere,andthe oceans are actlvely controlled heterotrophs Organismsthatobtainnourishment
by life, for life. from the tissues of other organisms.

geneticvariabilityA measure of thenumber of homeostasisAsteady-statewithinputscounter-

genetic diversity wlthin a gene pool. balancing outputs.

genus (plural genera) A taxonomic group of lower homeotherm An animalwhichregulatesits body

rank thana family that consists of closely related
specles or, in extreme cases, only one specles.
geodiversity The diversity of the physical environ-
ment.
geographical range Thearea occupied by a group of
organisms (species, genus, family, etc.); the same as
the distribution.
geosphere The solid Earth - core, mantle,and
crust.
geothermal springs Springs of water made hot by
the Earth’s internal heat.
germination The time of sprouting in plants.
G o n d w a n a Ahypothetical, Late Palaeozoic super-
continent lying chiefly in the Southern Hemisphere
temperature by mechanisms withln its own body; an
endotherm or ‘warm-blooded’animal (cf. poikilo-
therm).
homogeneousReferringtoa mosaic of similar
spatial elements; uniform.
humification The process of humus formatlon.
hummock-hollow cycle A peatland cycle involving
the infilling of a wet hollow by bog mosses until a
hummock is formed and colonized by heather. The
heather eventually degenerates and the process starts
again.
h u m u s An amorphous, colloidal substance produced
by soil micro-organisms transforming plant litter.
hurricaneAviolentandsometimesdevastating
tropical cyclone.
 GLOSSARY 23 1

hydrophyte A plantthatgrows wholly or partly LaurasiaTheancient,NorthernHemisphereland

submerged in water. mass that broke away fromGondwanaabout 180
million years ago and subsequently split into North
hydrosphere All the waters of the Earth. America, Greenland, Europe, and Northern Asia.
hyperparasite A
parasite that lives on another
leaf-areaindexTheratio of total leaf surface to
parasite.
ground surface. For example, a leaf-area index of 2
hyperthermophile An organismthatthrives in would mean that if you were to clip all the leaves
ultra-hot conditions. hanging over 1 m2 of ground, you would have 2 m2
of leaf surface.
Ice Age An oldterm for theQuaternary glaclal-
interglaclal sequence. legume A plant of the family Leguminosae, charac-
teristically bearingpodsthatsplitintwoto reveal
iceage A timewhen Ice formsbroad sheetsin
seeds attached to one of the halves.
middle and high latitudes (often in conjunction with
the widespread occurrence of sea ice and permafrost) lichen A plant consisting of a fungus and an alga.
and mountain glaciers form at all latitudes.
lifetable A tabulation of thecompletemortality
interspecific
competition
Competition
among schedule of a population.
species.
life-form The characteristics of a mature animal or
intestinal
endosymbiontsSymbloticorganisms plant.
living in another organism's gut.
littoral Shallow water zone
intraspecificcompetitionCompetitionwithin a
local extinction (extirpation) The loss of a species
species.
from a particular area.
intrazonal soils Soils whose formation is dominated
macronutrients Chemical elements needed in large
by local factors of relief or substrate; an example is
quantltles by living things.
soils formed on limestone.
introducedspecies or introduction A species maquis A shrubbyvegetation of evergreen small
released outslde Its natural geographlcal range. trees and bushes.

isolines A lineon a mapjoiningpoints of equal mass extinctions Extinction episodes in which large
value. numbers of specles disappear.

Jurassic period A slice of geological time spanning
208 million to 145.6 million years ago; the middle
unit of the Mesozolc era.
keystone species A specles that plays a central role
ina communlty or ecosystem, its removal havlng
far-reachlng effects.
landbridge A dry land connection between two
land masses that were prevlously separated by the sea.
land ethic A set of ethlcal princlples declaring the
rights of all naturalobjects(animals,plants,land-
scapes) to continued exlstence and, In some places at
least, continued exlstence In a natural state.
meadow Grassland mown for hay.
mechanistsProponents of the vlew thatall bio-
logical and ecological phenomena may be explained
by the interaction of physlcal entities.
megafauna Large mammals, such as mammoths and
sloths,contrastedwithsmallmammals, such as
rodents and insectivores.
megaherbivores Large browsers and grazers, such as
elephants and rhinoceroses.
mesophyte A plant flourishing under mesic (not too
dry and not too wet) conditions.
23 2 GLOSSARY
Mesozoic era A slice of geological time spanning
245 million to 65 million years ago comprising the
Triassic, Jurassic, and Cretaceous perlods.
metabolic processes The many physical and chem-
ical process involved in the maintenance of life.
micronutrients
Chemicalelements required
small quantities by at least some livlng things.
micro-organism An organismnot visible tothe
unaided naked eye.
mineralization The release of nutrients fromdead
organlc matter.
Miocene epoch A slice of geological tlme spanning
23.3 million to 5.2 million years ago.
mutualism An obligatoryinteraction between two
species where both benefit.
natural selection Theprocess by which the environ-
mental factors sortand sift geneticvariabilityand
drive evolution.
Neogene period A slice of geological t m e between
23.3 million and 1.64 million years ago.
net primary production Gross primary production
less respiration.
neutralism An lnteractlon in whlch neither species
is harmed or benefited.
neutrophiles Plants that like neutral conditlons, not
too acldic and not too alkaline.
nitrogen-fixingThe conversion of atmospheric
nitrogen
into
ammonium compounds by some
bacteria.
nutrientA chemical element required by at least nity in marlneand freshwatersystemwhich
some living things.
old-growthforestAvlrgln forest that has never and diatoms.
been cut, or a forest that has been undisturbed for a
long tlme.
Oligocene epochA slice of geological time spanning
35.4 million to 23.3 million years ago; the youngest iognomy (life-form) occupying a climatically uniform
unit of the Palaeogene period.
orobiomes Biomesassociated withthealtitudinal
climatic zones on mountains.
overwinterTo spendthewinter in aparticular
place.
Palaeocene epoch A slice of geological time span-
in ning 65 million to 56.5 million years ago; the oldest
unit of the Palaeogene period.
PalaeogeneperiodA slice of geological t m e
between 6 5 million and 23.3 million years ago.
Palaeozoic era A slice of geological time spanning
570 million to 245 million years ago; comprises
the
Cambrian,
Ordovician, Silurian, Devonian,
Carboniferous, and Permian periods.
Pangaea The Triassic supercontlnent comprising all
the present continents.
pathogen Anagent, such as bacterlum or fungus,
that causes a disease.
pedobiomes Areas of characteristic
vegetation
produced by distinctive soils.
pedosphere All the soils of the Earth.
Permian period A slice of geological time spanning
290 million to 245 million years ago.
photoperiod The duration of darkness and light.
photosynthesis The synthesisof carbohydrates from
carbon dioxide and water by chlorophyll, using light
as an energy source. Oxygen is a by-product.
phytomass The mass of living plant materlal in a
community, or in a unit area; usually expressed as a
dry welght.
phytoplanktonPlantplankton:theplantcommu-
floats
free in the water and contains many species of algae
phytotoxic Poisonous to plants.
plantformationThevegetatlonalequivalent of a
biome, that IS, a community of plants of like phys-
area on a continental scale.

Pleistocene epoch A slice of geological tune span-
ning 1.64 million to 10,000 years ago; the older unlt
of the Plelstogene period (or Quaternary sub-era).
Pleistogene period The mostrecentslice of geo-
logical time between 1.64 million years ago and the
present;subdividedinto Pleistocene and Holocene
epochs.
Pliocene epoch A slice of geological time spanning
5.2 millionto 1.44 million years ago;theyounger
unit of the Neogene period.
poikilotherm An organism whose body temperature
is determined by the ambient temperature and who
can controlits body temperature only by taking
advantage of sunandshadeto heat up or to cool
down; a 'cold-blooded' animal (cf. homeotherm).
prairie An extensive area of natural, dry grassland;
equlvalent to steppe In Eurasla.
profundal Deep water zone.
propagule The reproductive body of a plant.
protocooperation
non-obligatory
A interaction
between two species where both benefit.
psychrophile
Organisms
that flourish at low
temperatures.
race An anlmal or plant populatlon that differs from
other populations of the same specles in one or more
hereditary characters.
radioisotopeA form of a chemical elementthat
undergoes spontaneous radioactlve decay.
rain shadow A region wlth relatively low rainfall
that is sheltered from raln-bearingwinds by high
ground.
respiration A complexserles of chemical reactions in
all organlsms by which energy is made available for
use. The end products are carbon dioxlde, water, and
energy.
rhizome A root-like stem, usually horizontal, grow-
ing underground with roots emerging from Its lower
surface and leaves or shoots from Its upper surface.
GLOSSARY 233
salinity Saltiness.
saltatorial Adapted for hopping and jumping.
savannah Tropical grassland.
saxicolous Growlng on or living among rocks.
scavenger An organism feedingon dead animal flesh
or other decaying organic material.
seasonality The degree of climatlc contrast between
summer and winter.
sessile
Permanently
attached or fixed; not free-
moving.
social Darwinism Darwln's doctrme of the survlval
of the fittest applied to society.
soil Rock at, or near, the land surface that has been
transformed by the blosphere.
solar radiation The total electromagnetlc radiation
emltted by the Sun.
Electromagneticradiatlon
is energypropagatedthrough space orthrougha
material as an interaction between electric and mag-
netic waves; occurs at frequencies rangmg from short
wavelength,high frequencycosmlc rays, through
medium wavelength,
medium frequency
vlsible
light, to long wavelength,low frequency radio waves.
speciation The process of species multlplication.
species A reproductively isolated collection of inter-
breeding populations (demes).
steppe An extensive area of natural, dry grassland;
equlvalent to prairiein North American terminology.
stomata (singular stoma) Epidermal pores in a leaf
or stem through which air and water vapour pass.
sweepstakes route A path along whlch organisms
may disperse, but very few will do so owing to the
difficulties involved.
symbiotic Pertaining to two or more organisms of
differentspecies livlngtogether.Ina narrow sense
equivalent to mutualism.
taxon (plural taxa) A population or group of popu-
lations (taxonomic group) that is distinct enough to
234 GLOSSARY

be given a distinguishing name and to be ranked in ungulates Hoofed mammals.

a definite category.
vascular plant A plant containing vascular tissue,
territoryAn area defended by an animalagainst theconducting system which enables waterand
other members of its specles (and occasionally other minerals to move through it.
species).
vegetation Plant life collective; the plants of an area.
Tertiary period A slice of geological time spanning
vernalization The exposure of some plants (or their
65 million to 1.64 million years ago and the youngest
seeds) to a period of cold that allows them to flower
unit of the Cenozoic era; now designated a sub-era.
or to flower earlier than usual.
theory of islandbiogeography A modelthat
vicariance The splitting apart of a species distribu-
explains the species diverslty of Islands chiefly as a
tion by a geological or environmental event.
function of distance from themainlandand island
area. vicars Species of different ancestry but with the same
characteristlcs living indifferent
biogeographical
thermophiles Organisms that love hot conditions.
regions. Also called ecological equivalents.
toposequence The sequence of soils and vegetation
vitalists Proponents of the Idea that some kind of
found along hillslope,
a from summitto valley
living force resides in organisms.
bottom.
volatilization To convert to vapour.
trace elementA chemical element used by an organ-
ism In minute quantities and vital to its physiology. wetlands All the places in the world that are wet for
at least part of the year and support a characterlstic
tree-lines The altitudinal and latitudinal limits of soils and vegetation; they Include marshes, swamps,
tree growth. and bogs.
tree-throw The toppling of trees by strong winds. xerophyte A plant adapted for life in a dry climate.
Triassic period A slice of geological time spanning zonobiome One of nine, climatically defined, major
245 million to 208 million years ago; the oldest unit community units of the Earth.
of the Mesozoic era.
zoomass The mass of livinganimalmaterialin a
trophic
Of, or pertaining
to,
nourishment or community, or in a unit area; usually expressed as a
feeding. dry weight.
tsunamis The scientific (and Japanese) name for tidal zooplanktonAnimal
plankton;
aquatic inver-
waves. tebrates living in sunlit waters of the hydrosphere.
 FURTHER READING

General
biogeography
books Borhldi, A. (199 1) Phytogeography and Vegetatton
Ecology of Cuba, Budapest: A k a d h l a l Kiad6.
Cox, C. B. and Moore, P. D. (1993) Biogeography A n
Ecologtcal and
Euoluttonaty
Approach, 5thedn,Oxford:Bowman, R , I, (ed,) (19(;6) The Gala'pagos
Blackwell. (Proceedings of theSymposium of theGalipagos
Already a classic. A very popular textbook. International Sclentlfic Project), Berkeley and Los
George, w, (1962) Anitna(Geography, London:Angeles,California: University of Californla Press.
Helnemann.
Written before the revival of continental
drift,
H.C. (ed.) (1984) and BiogeograPhY
well
but
worth a look. in Sri L n k a (Monographiae Biologicae,
Vol. 57), The
Hague: W. Junk.
Pears, N. (1985) Basic Btogeography, 2nd
edn,
Harlow:
Longman.
Gressitt,
J. L.(1982)
(ed.) Biogeography and Ecology of
A good textbookonecologicalbiogeography. New Guinea, 2 vols (Monographiae Biologicae,Vol.
42), The Hague: W. Junk.
Tivy, J. (1992) Biogeography: A Study o f Plants tn the
Ecosphere, 3rd edn, Edinburgh: Oliver & Boyd.
Kuschel, G . (ed.) (1975) Biogeography and Ecology in
A good introductory textbook.
Neu, Zealand (Monographlae Biologicae,Vol. 27),
The Hague: W. Junk.
Regional biogeography
Stoddart, D. R. (ed.) (1984) Biogeography and Ecology
Here is a small selection of regional biogeographlcal
ofthe Seychelles Zslands (Monographiae Biologicae, Vol.
and ecological texts. A GEOBASE search will reveal
5 5 ) , The Hague: W . J u n k .
many more.

Battistini, R. and Richard-Vindard, G . (eds) (1972) Wallace's Line and Plate

Whitmore, T. C. (ed.) (1981)
Bzogeography and Ecology in Madagascar (Mono- Tectontcs (OxfordMonographs inBiology, Vol. l),
graphiae Biologicae, Vol. 21), The Hague: W . J u n k . Oxford: Clarendon Press.
 236 F U R T H E RR E A D I N G

Whitmore, T. C. (ed.)(1987)BiogeographicalEvolutron Whelan, R. J. (1995)The Ecology of Fire, Cambridge:

of the Malay Archipelago (Oxford Monographs in Cambridge University Press.
Biology, Vol. 4), Oxford: Clarendon Press. A clear introduction to the subject.

Williams, W.D. (ed.) (1974) Biogeography and

Historical aspects
Ecology zn Tasnrunta (Monographlae Biologicae, Vol.
Hague:
The25), Junk.
W. Brlggs, J.
(1995)
C. Global Bzogeography
(Developments In Palaeontology andStratigraphy,
Woods,C. A. (ed.)(1989) Bzogeography of the We.rt 14)~Amsterdam: Elsevier.
Indies: Past. Present. and Fzrtzrre, Gainesville,Florida: An advanced text but worth dipping into.
Sandhill Crane Press.
Cox, C. B. and Moore, P. D. (1991) Bzogeogruphy: A n
Erological and Evolutionary Approach, 5th edn, Oxford:
aspects Ecological Blackwell.
Dansereau,
(1957)
p. Biogeography: An Eco/ogll~/ basic coverageOf historical aspects.
Perspective, New York: Ronald Press.
Cronk, Q. C. B. and Fuller, J. L. (1995) Plant
Old but have a look.
Invaders: The Threat t o Natural Ecosystons, London:
Chapman & Hall.
Forman, R. T. T. (1995) L n d Mosaics: The Ecology o f
Many examples.
hzdsrapes
and Regions, Cambridge:
Cambridge
University Press.
Darlington,
P.
J.,
Jr
(1957) Zoogeography: The
A weighty tome on landscape ecology with some
Geographical Dzstributron o f Aninfal.r, New York: John
useful sections for biogeographers.
Wiley & Sons.

Kormondy, E. J. (1996) Concepts of Ecology, 4th edn, and Plants, London: Chapman & Hall.
Englewood Cliffs, NJ: Prentice Hall. A little gem.
An excellent basic ecology text.
Hallam,
A.
(1995) A n Ozrtline of Phanerozoic.
Biogeography, Oxford: Oxford University Press.
Larcher, W. (1995) Physzologtcal Plant Ecology:
Ecophyszology and Stress Phystology of Functional Groups, A palaeontological perspective.
3rd edn, Berlin: Springer.
First-ratecoverageofphysiologicalaspectsof Nelson, G.and Rosen, D. E. (eds) Vicariance
plant ecology. Biogeugraphy: A Crrtrqzre (Symposium of the
Systematics
Discusslon Group of the American
Stoutjesdijk.
and
Barkman,
P. (1992)
J.J. Museum of Natural History, 2 4 May 1979), New
Microdimate, Vegetatron and Furma, Uppsala, Sweden: York: Columbla University Press.
Opulus Press. Only for the brave.
Unusual but highly interesting book.
Udvardy, M. D. F. (1969) Dynumtc Zoogeography, u'rth
Viles, H. A.(ed.)(1988) Brogeonlorphology, Oxford: SpeL'ialReference to LandAnzmah, New York:Van
Basil Blackwell. Nostrand Remhold.
Links life to geomorphology. Rather old, but still deserves to be read.
 FURTHER READING 237

Wallace, A. R. (1876) The Geographrcal Distrihutron of Grover, J. P. (1997) Raource Competitron, New York:
Animals: with a Study of the Relattons of Ltvrng and Chapman & Hall.
Extrnct Faunas asEluc1datzng the Past Changes of the Up-to-date exposition.
Earth's Suvface, 2 vols, London: Macmillan.
All biogeographers should read this great work. Jackson, A. R. W. and Jackson,J. M. (1996) Environ-
mental Science: TheNatural Environment andHuman
Woods,C.A.(ed.) Biogeography of the West Indies: Impact, Harlow: Longman.
Past. Present. and
Future, Gamesville, Florida: Excellent text.
Sandhill Crane Press.
Caribbean case studies.
Kingsland, S. E. (1985) ModelingNature: Eprsodes rn
the History of Population Brology, Chicago and London:
Single populations Unlverslty of Chicago Press.
Hanskl, I. and Gilpin, M. E. (1997) Metapopulation A first-rate account of the history of population
Biology: Erology,Genetrcs. and Evolution, New York: biology. More exciting than it might sound.
Academic Press.
Advanced but worth reading. Kormondy, E. J. (1996) Concepts o f Ecology, 4th edn,
Englewood Cliffs, NJ: Prentice Hall.
Kormondy, E. J. (1996) Concepts of Ecology, 4th edn, Clear explanations of basic ideas.
Englewood Cliffs, NJ: Prentlce Hall.
Relevant sections worth reading. MacDonald, D. (1992) The Velvet Clauj: ANatural
History ofthe Carnivores, London: BBC Books.
Kruuk, H. (1 995)W i l d Otters: Predatzon and Poprda-
All about carnivores with beautiful photographs.
Oxford: Oxford University Press.
trons,
A good case study.
Communities
Perry, J.H.,Wolwod, I. P.,Smith,R.H.,and
Morse, D. (1997) Chaos in Real Data: Analysis on Non- Archibold, 0. W. (1994) Ecology o f World Vegetation,
lineur Dynanmlr from Short Ecologiczl Time Serres, New York: Chapman & Hall.
London: Chapman & Hall. Covers ecological aspects of plant communities.
If mathematics is not your strong point, forgetit.
Chameides, W . L. and Perdue, E. M.
(1997)
Taylor, V. J.andDunstone,N.(eds) (1996) The Cycles: A Cot~I~uter-lntwactiveStudy o f
Bmgeoo'hen~~czl
Exploitation o f M a m d Populations, London: Chapman Earth System Sc1enr.c andGlobalChange, NewYork:
& Hall. Oxford University Press.
Examples of exploitation. If you demand much, this is the one for you.

Tuljapurkar, S. and Caswell, H.(1997) Structured-

Hochberg, M. E., Colbert, J., and Barbault, R. (eds)
Populatlon Models, New York: Chapman & Hall.
(1996) Aspects of the Genesrs and Marntenance o f
Somewhat demanding.
Biological Dtversity, Oxford: Oxford Unlversity Press.
Contains many case studies, though not an easy
Interacting populations read.
Crawley, M. J.(1981) Herbluory: TheDynamirs of
Anmal-Plant Interac-tions (Studies in Ecology, Vol. Jeffrles, M. L. (1997) Blodiversltyand Conservation,
lo), Oxford: Blackwell Scientific Publications. London and New York: Routledge.
A
superbbook,
despitethe
mathematics. An
excellentbasictext.
23 8 F U R T H E RREADING

Lawton, J. H. and May, R. M. (eds) (1995) Extinctton Community change

Rates, Oxford: Oxford University Press.
Lots of recent figures on extinction rates. Committee
Characterizatlon
on of Wetlands,
National Research Council (1995) Wetlands:
Morgan, S, (1995) El.ology and
Envtronment:
The cycles and Boundaries* Washington, DC:
of Lge, Oxford: Oxford University Press. National Academy Press.
A good starting text. Readable case study.

Polis, G . A.andWinemiller, K. 0. (1995) Food Gates, D. M. (1993) Climate Change and Its Btologrcal
Webs: Integration ofparternsandDynamtlJ, New York: Consequences, Sunderland, Massachusetts:Sinauer
Chapman & Hall. Associates.
Not easy, but worth a look. A clear account.

Quammen, D. (1996) The Song of the Dodo: Island Huggett, R. J. (1993) Modelling the Human Impact on
Biogeography in
an Age of Extrnctions, London: Nature: Systems Analysis of Environmental Problems,
Hutchinson. Oxford: Oxford University Press.
Highly readable. If youlikemodellingbutarenottoomathe-
matically minded, this might be of interest.
Reaka-Kudlka, M. L., Wilson, D. E., and Wilson, E.
0. (1997) Bzodiversity 11: Understanding and Protecting Huggett,R.J.(1997) EnvtronmentalChange:The
Evolving Ecosphere, London: Routledge.
Our Biological Resources, Washington,DC:National
Academy Press. Provides a broad perspective.
Sure to become a classic.
Jackson,
A. R. W . andJackson,J.M.(1996)
Rosenzweig, M. L. (1 995) Speczes Dzversity zn Space and Environmental Science: TheNatural Enzsronment and
Time, Cambridge: Cambrldge Universlty Press. Human Impact, Harlow: Longman.
Fascinating, but not easy. A basic textbook with several relevant sections.

Schultz,J.(1995) The Ecozones of the World:The Matthews, J . A. (1992) The Ecology of Recently-
Ecological Dtvzsions of the Geosphere, Hamburg: Deglaciated Terrain: A Geoecologrcal Approarh to Glacier
Springer. Forelands and Prtmnary SuccesJton, Cambridge:
A verygoodsummary of theworld’smain Cambridge Universlty Press.
ecosystems. An excellent case study and lots more.

Szaro, R. C. and Johnston, D. W. (1996) Biodiversity Peters, R. L. and Love~oy, T. E. (eds) (1992) Global
Warmingand
Biolopcal
Dtuersity, New
Haven,
in Managed Landscapes: Theory and Practice, New York:
Connecticut and London: Yale University Press.
Oxford Untversity Press.
Try it. A host of examples in this one.

Schwartz, M. (1997)
Conservation z n Highly Fragmented
Wilson, E. 0. (1992) The Diversity of Life,
Cambrldge, Massachusetts: Belknap Press of Harvard LndslilpeJ’ New Chapman &
Topical and highly recommended.
University Press.
A highly readable book.
 F U R T H E RREADING 239

Ethical issues and conservation

This is a selection from a long list.

Anderson, E. N . (1996) Ecologies of the Heart: Emotion,

Belid and the Environment, New York: Oxford
Unlversity Press.

Botkin, D. B. (1990) Discordant Harmonies: A Neuj

Ecology for the Twenty-First
Century, NewYork:
Oxford University Press.

Committee onScientific Issues in theEndangered

Species Act, National Research Council (1995) Science
and the Endangered Specres Act, Washington,DC:
National Academy Press.

Cooper, N. andCarling, R. C. J. (eds) (1996)

Ecologrsts and Ethical Judgements,London: Chapman &
Hall.

Norton, B. G. (1991) Toward Unity Among

Envzronmentalists, New
York:
Oxford
Universlty
Press.

Ryder, R. D.(ed.) (1993) AnimalWeyareand the

Environment, London: Duckworth and RSPCA.

Shafer, C. L. (1990) Nature Reserves: Island Theovy and

Consenlation Practice, Washington,DCand London:
Smithsonian Institution Press.

Sylvan, R. andBennett,D. (1994) The Greenrng of

Ethrcs: From Human Chauvinism t o Deep-Green Theory,
Cambridge: White Horse Press; Tucson: University
of Arizona Press.
 BIBLIOGRAPHY

Ables, E. D.(1969)‘Homerangestudies ofredfoxes Baker, A. J . M., Proctor, J., and Reeves, R. D. (eds) (1992)
(Vrdpes vrdpes)’, Journalof Manrntalogy 50: 108-20. The Vegetatron o/ Ultranrajc(Swpentme) Soils, Andover,
Agee. K.
J.
andJohnson,
D.
R.
(1988) Ecosystem Hants: Intercept.
Management for Parks and Wilderness, Seattle, Barfield, C. S. and Stimac, J. L. (1980) ‘Pest management:
Washington: University of Washington Press. an entomologlcal perspective’, BioScrence 30: 683-8.
Allen, R. (1980) How t o Save the World: Strategy /or World Bartholomew, G . A. (1968) ‘Body temperature and energy
Consmatron, Foreword by Sir Peter
Scott,
London: metabolism’,pp.290-354inM. S. Gordon, G. A.
Kogan Page for IUCN, UNEP, and WWF. Bartholomew, A. D. Grmnell, C. B. Jorgensen, and F.
Anderson, P. and Shimwell, D. W. (1981)Wild Flouws and NW. h l t e (eds) Anrmal Functron:Prrncrples and
otherPlants of thePeak District: A n Ecologrcal Study, Adrptafions, New York: Macmillan.
Ashbourne, Derbyshire: Moorland Publishing. Bartlett, D. H. (1992) ‘Microbiallife a t high pressures’,
Arnold, G. W . , Steven, D. E., Weeldenburg, J . R.,and ScrenreProgress 16: 479-96.
Smith, E. A. (1993) ‘Influences of remnant sue, spacing Bascompre, J. andSole, R. V. (1995) ‘Rethinkmg com-
pattern and connectwlty on population boundaries and plexlty: modelling spatiotemporal dynamics in ecology’,
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 INDEX

aardvark (Ovt-terOpuru/er), African ant-eater 127 Arctic avens ( D ~ urntegr@iu),

J heliotrope 18
abiotic environment 4 Arctic fox (Alopex logopur). adaptation to cold 20
acclimatization 22, 210 Arctic poppy (Puparm mdimtuni), heliotrope 18
accommdationists (environmental managers) 21 3 area-alone hypothesis, in island biogeography I63
acidophiles 30 armyworm (Spodopteru exemptu), r-strategist 99
acorn barnacles (Chthumufurrteffuturand BulunuJ hafunorder), aspect, as environmental factor 32
aggressive competitors 116 austral (southern) distribution ree distribution
active competition 7 6 autwcological accounts 37, 39, 42
aerial, adaptation i n mammals 37 autogenic succession ree succession
aggressive competition 1 1 6 available moisture 2 3
alkaliphiles 30 azuki bean weevil (Cu//orobrrd~~b~ds
chrnenrts), prey species in
allogenic succession ree succession laboratory experiments 132
alpine belt, on mountains 30
alpine marmot (Alomrotu ntuv~rotu):and aspect 33; as climatic background matrixes 14
relict 49 balanced ecosystem(s) 172, 179, 208, 219, 221-3
alpine snow buttercup ( R u ~ ~ ~ L .udoneur),
u ~ I L J heliotrope 1 8 BUlUnUr bu/unoider, aggressive competitor 1 16
altitude, as environmental factor 32 banner-tailed kangaroo rat (Dipodon~yrJpertu6dir). water intake 26
Amargosa vole (Mirrotur culi/iornrcrdr ri-rrpenrir), micro-endemic 43 barophiles 15
amensalism 120 barrel cactus (FWJUCIUJ u'rrlizetri), a succulent 27
American bison or 'buffalo' (Biron brron), overexploited species barriers (to dispersal) 61, 7 0
105 Batesian mimicry 112
American chestnut ( C U J ~ ~ T&ntutu)
IU 80, 156, 203 bay bamboo-rat ( C ~ n n o n t 6udiu.r),
y~ prey species in southrrn India
American mink (Murrefuvrsotr), in Britam 78 128
American muskrat (OnLrru zibetbrrur), in Europe 5 8 beaver (Curtor runudenrrr),and commensalism 113; keystone
American pronghorn (Antilocopra uvierr~imu), adaptation to cold herbivore 154
20 Belding ground squirrels (~pernrophi/uJbdditrgi), adaptation to
American robin (TurduJ nirgruroriur ~ I ~ U ~ O Y Isurvivorship
~ J ) , short summers 18
curve 92; in fruic-frugivorc system 124 big sagebrush (Arrenlrrru rridevtutu), true xerophyre 27; in
amphibians. and global warming 198 Nevadan ghost towns 178
amphitropical distribution ree distribution bighorn sheep (OUIJ spp.), avoiding warble flies 27
analytical biogeography 1, 4 biochore 59
anemochore 59-60 bioclimates 23, 40
anemohydrnchore 59 biocwnoses 138
animal liberation movement 210 biodiversity 142, 159, 1 6 6 9 , 197, 212, 219, 224
animal life-forms 37 biogeochemical cycles 149-50
arboreal, adaptation in mammals 37. 156 biogenchemicals 149, 169
 INDEX 255
chuckwalla (.Y'iiiroiiidm ubemr), water loss 25
CITES (Convention on Inrernaciimal Trade i n Endangered
S ~ x c i r sof Wild Fauna and Flora) 2 1 I
classiticariim 6
climate diagrams 23-1
climatic climax (vegetatinn) 171-2, 173. 179, 208
climatic disjunctions 4X
climatic relicts .?9
c l i m a t i c a l l y sensirive qwcies 197, 208
climax-pattern hypothesis 172
1 Ioudrd Icopard (Nef&'/ii NPbiikII~i).predator in Thailand I 2 8
Coastal Eci)logical Landscape S p a t i a l Simulatiim rn(xlt.1 (CELSS
mudel) 191
cixistal ecosystem. southern Liiuisiana I9 I
cocklebur (S,rii/biiiiri J/riiiridriiim), short-day plant I X
ciihi)rt-survival mixlels 92, I O X
ciild-ti)lrrant plants 2 2
wlliired lemming (DiitDitiiii)x spp.). individualistic response to
climntic change 1x2
ionima butterfly (Pd)fioiii(i i--&riiii), response to climatic
warming 197
ciimmensal(s) 109, I 1 3 , 128, lZ6
c~inimensalism 109. 11.5, 136
cimmon p y ( G m d t i t xl~riiilmwr),inhahirant of deciduuus forest
15
cummiin myna (c\<rrclorheriil ~ r i t t i ~and ) . commensalism I 1 i
cummiin i d ( Q i i c r w robiir), unpalat;ihlc lcaves id I 2 3
c i m m o n wallaron or eiirii (AI'r[-ropiil robiirtiii). i n fragmented
landscape 9 X
common water crowfoot (KdiiroiliiIi/r q i ~ ~ r d), ihydropliyre i 24
communalists (self-reliance. soft technoliigisrs) 2 15
communities, dctined 1%; i n the twenty-tirsr century 197;
impermanence 179. 1 X 2
compensation level 143
competitive exclusion principle 36, 114, 136
<ompetitive ruderals 1 0 2
competiti,r-stress-rolerator-ru~lrral ( G S - R ) 1 0 i
ci)mtxritors-cum-insinuators7 6
(:i)mpiisitae. widespread plant family 44
consumers (heterimophs) 142
consumption zone 1.12-.3
coiiresr (interference) competition 114. 116, 1 56
ciintinental drift 8 . 67, 7 0 , 7 2
continental tission 67
cimtinrntal ~ U S I O I I7 6
contincntaliry 18
continuiiiis ilistrihurion JCP disrrihurion
Cuiiper's hawk (/\c.t ipitcr iooprri), ;ind rcsiiiircc partitioning
I17
ci)priiphages 1I 5
c~irnuciipians(iiprimists) 2 I i
ci)smqxilit;in species (cosmopolites) 43
cnypii ( A l ) a . r r t o r c u p i J ) , i n Hritain 77
criissulacean acid mctahnlism (CAM) 27
creiisi~tehush ( L i i w r d i ~ , ~ i r i w dri)ught/~~), rndurrr 27
Croizar. LCon, f'ither (if viciiriance biogeography 67
cryptophyre .i7
culling IOX. 210-11, 2 1 2 . 22.1
cursimal. ;idapt.ition in mammals 17, 72
cycads 5 0
256 INDEX

Ihll m o u n t a i n sheep ( O w Jdlr JuNr), s u r v i v o r s h i p c u r v e 9 2 e m e r g e n t properties (of c o m m u n i r i e s ) 1 2 , I85

d a n a i d b u t t e r t l i e s (Danainae). as mwlels in m i m i c r y cnmplexes e m i g r a t i n n ( a n d i m m i g r a t i o n ) 84
I23 emos s k i n k s ( l h r u nrgru a n d h o r u m w d u ) ~ and inrrciluced
clandelion (Turuwt-rim o/$citiu/e), f u g i t i v e - m i g r a t i o n strategist 1 0 3 predators 80
D a r w i n . Charles R o b e r t 63, 67. 212, 2 1 5 . 223 e n d e m i c species 32, 4 3 , 1 6 5
day-degrees 22, 55. 56 e n v i r o n m e n t a l economics 222
DDT 158-9, 2 I 3 e n v i r n n m e n t a l ethics 209
decomlwser ( d e t r i t n l ) fi~d web I 5 1 e n v i r u n m e n t a l i s m 209, 2 13-15, 223-4
decompcrsers (sapriiphages) 142. 117-8, 15 1, 169 E s k i m o dog, a d a p t a t i o n tn c o l d 2 0
~ l e c o m p o s i t i u n148. 149 e u r o or c n m m o n w a l l a n x ) (Afurropm robrtstrdr), in f r a g m e n t e d
deep ecologists (Gaians) 2 I5 landscape 98
deep ecology 216 European badger ( A f e b d e r ) , a n d p r e y s w i t c h i n g 128
1)egeneriact.ac. restricted endemic p l a n t f a m i l y 44 E u r q m n beaver (C'uJ/urfiber)3r e i n t r c x l u c t i o n in Sweden I06
demes 84, IO X European n u t h a t c h (Sirtu europueu), as m e t a p i p u l a t i n n 95
desert night l i z a r d (Xun/urtu U ~ I / J J ) , s u r v i v i i r s h i p curve 92 , fwal source for badger
European r a b b i t (Oryitolugris m i i r w / r ~ . r )as
designer mosaic 14 I28
d e t r i t a l (decomposer) fwxf web I 5 I E u r o p a n s t a r l i n g (S/iwint.r d p r r i l ) , dispersal in North A m e r i c a
d c t r i r i v o r e s (saprovores) 121. 1.42. 147-8. 151, 1 6 9 63
D e v i l ' s Hole p u p f i s h (Cjprrm)dnir druboh), m i c r o - e n d e m i c 4 3 Everglade (or snail) k i t e (/h/rhunirdJ Jocruhr/r.r), h i g h l y spxialized
clhole (Caotr u/piiiiii), predatnr in s ~ i i i t h e r n1ndi;i 128 S[X"'S 197
D I ~ I N I NpIn N i t o. z n a n predatnr in laboratory experiments 130-2 evolutionary diseqiiilibriuni 223
d i f f i i s i n n (of species) 5 8 , 1.$7 evoluririnary d i s ~ u n c r i ( i n s46
d i s c l i m a x ( p l a g i n c l i n i a x ) 171. 17') e v n l u t i o n a r y relicts 49-50
d i s e q u i l i b r i u m : c n m m i i n i t i e s 171, 179. 219; ecologists 223; exponential g r o w t h 84, 86. 8 7
v i e w 179, 1 8 3 extinction-and-col[inization m e t a p o p u l a t i o n 95-6
d i s h a r m o n i o u s c o m r n ~ n i t i e s ,3, 184-5 e x t r r m i i p h i l e s 15
d i s j u n c t ( b r o k e n ) ~ l i s t r i b u t i o nSL'C d i s t r i b u t i o n ezo- m a m e - y anag i (Sulrx puucr/loru), acc I im a r izat ion in 2 2
disprsd 5; a b i l i t i e s 60, 197; routes 8. 61. 67, 8 2
d i s t r i b u t i o n : a n i p h i t r o p i c a l 45-7; austral (southern) '45; h ) r e a l f a c i l i t a t i o n m o d e l (of succession) 1 7 3 4 , 208
( n o r t h e r n ) .45; b r o k e n ( d i s j u n c t ) 46, 69; c n n t i i i u o u s 4 6 ; false ragweed (Pur/herrrii?//hy/erophor/ds),b i o l r i g i c a l c(inrrol of I35
p a n t r i i p i c a l 4 5 ; tempcrate ( m i d d l e latitude) 4 5 faunal m i x i n g 76, 83
disturbance. as enviriinment;il factor 33-5; a n d forest dyn:imics faunal strata 66
206; b y e p i d e m i c s 89 f e c u n d i t y 8.4. 88, 90. 91, 93, 94, 97, 99
d i v e r s i t y : change 166; hot-spots 165, 169; l a t i t u d i n a l g r a d i e n t f e r t i l i t y 84
161, 163-6, 169; rep u/w b i o d i v e r s i t y f i l t e r routes 62
d o c k (l</imi,x iihtimfulrns), in horse pasriires I 2 5 fire, as ;in e n v i r n n m e n t a l factor 35
d o m e s t i c cats (FelrJ ~ i t t i i i )on , M a r i o n I s l a n d 107 I ~ l o r i d ap o i s o n tree (hlerriprr~rntoxifirm), p r e d i c t e d d i s t r i b u r i o n
cloiibling t i m e 86 56
d u c k w e e d (1.emu) species. c o m p e t i t i o n among 1 I(> fmxl webs 142, 351-.3. 169. 185; c o n i a m i n a t i o n of 156
dwarf shrew (Snrex m i w ~ )saxicnlnus , species iI foO&hain c o n c e n t r a t i o n ( b i n m a g n i f i c a t i o n ) 158
forest: change in eastern N o r t h A m e r i c a 201; disturbance 206;
eastern phoebe (Stiyormr phoebe), w i n t e r c l i s t r i b u t i i i n and g r o w t h m o d e l 190, 205; succession 2 0 6
abundance 56 f i i r m a t i o n - t y p e (zonal p l a n t f o r m a t i o n ) 3 0
e c o ~ e n r r i c215.~ 222 fossnrid ( b u r r u w i n g ) . adaptatinn in mammals 17
ecocentrism 213-15. 2 2 4 freshwater c o m m u n i r i e s 10
e c t i l i i g i ~ dhiogengraphy I , 4 frost-resistant plants 2 1
rcologic-al equivalents 16. 'I.?.77 frost-tolerant plants 2 I
ecnlngical n i c h e 35-6, 4 2 , 7 6 f r u i t - f r u g i v n r e system 12-1
cciilngical p y r x m i d s 15 I f i i g i t i v ~ - m i g r ~ i t i ostrategists
n IO i
ccnlngical tnlerance 15-17 f u n c t i o n a l : n i c h e 36; n u t r i e n t s 149
ecnlogic.il valency 16, 40 f u n d a m e n t a l ( v i r t u a l ) n i c h e 36
ecosphere .40, .12, 1.12. 1.49. 169, 173, 185, 2 2 2 f u n n e l a n t (ApburrnoguJtet./ i i q r ~ ~ e p topi)graphic
), intluencc o n
ecosystrm, defined 118; management LO?, 218-19, 2 2 3 . 22.4 d i s t r i b u t i o n 33
ecotones 2 8 , 10, 172, 201
ccoznnc's 27-9, 10, 36, 37. 40, 139 Gaia hypothesis 173, 221. 222
ELIitIi's checkerspur h u r r e r f l y (Eidphydty4.r edr/hu), a n d g l o b a l Gaians (deep ecologists) 2 I 5
w a r m i n g 198 gaps (in fiirests) 35
electrnm;ignetic r a d i a t i o n 17* 18 Gause's p r i n c i p l e 1 1 4
elephanr(s): African (Loxodorire &iiiuu) 88, 155; as s w i m m e r s 60 generic c o n t r o l 135-6
E/i,iith~'r,)d.n-r)//d, frogs, in Caribhean I i i s t n r i c a l biogeography 7 4 g e n e t i c d i v e r s i t y 159, 222
 INDEX 257

genus, defined 6 h o m e o t h e r m s 19-20, 26

geographically localized species 197 Homo .iupictr.i I 5 , 2 15
geological dislunctions 48 honey badger or ratel (hle//irarrr cupeiirrr), protocooperator 109
geomass 146 honey possum o r n o d b e n d e r (Trrrriper rortrrrtrrr), m a m m a l i a n
geosphere 149 p o l l i n a t o r 110
g i a n t or great anteater (Alyrtnemphrrp trrdul-tylu), adaptations for honey-guide (Indimtor riidrtwor), proroccx)perator I09
e a t i n g ants I27 house mouse ( A h nru.iiii/rrJ J O ~ N ~ aJn~ dK ~~J),
commensalism 1I3
glacial flea (Iroronru .ru/tunr), n a r r o w temperature tolerance of 16 housefly (Alum domerfnu).a n d exponential p o p u l a t i o n growrh 86
G l a n v i l l e f r i t i l l a r y (Ale/rtrrerr i.tnxru), m e t a p o p u l a t i o n in F i n l a n d H u a i Kha K h a e n g W i l d l i f e Sanctuary, T h a i l a n d 128
96 h u m m o c k - h o l l o w cycle 3.1
g l y p t c d n n t s , e x t i n c t t a n k l i k e armadillus from S o u t h A m e r i c a 65 h u m u s 148, 1/19
goats (Cuprrr hrrur), i n N e w Yxaland 107 hydrochore 59
g o l d e n l i o n t a m a r i n (Lroiitrdrrrr r o r a h ) , threatened species 197. hydrophytes 2354
212 hydrosere 174
g o l d e n toad (Bufo perr,q/ener), a n d glnbal w a r m i n g 198 h y p w r h e r m u p h i l e s 17, 19
G o n d w a n a 48,69-73
goshawk (Airiprter g e n f d r r ) . a n d resource p a r t i t i o n i n g I 17 Iceland purslane (Kurnrgru r.i/unhu),l o w w a r m t h requirement 22
Graminae, widespread p l a n t f a m i l y 44 i n c l i n a t i o n (slope), as environmenral factor 33
g r a z i n g food chain 1 5 1 , 170 I n d i a n mongoose (Ilerprmr iiiiropiitr'tirfrir), i n t r o d u c e d predator 7 9
G r e a t A m e r i c a n Inrerchange 67. 76-7. 83 inhibition m o d e l (of suc-cession) 174, 208
g r r a t a u k ( P J I I ~ ~ W Iinrpennrr),
UJ h u n t e d to e x t i n c t i o n 104 insinuators 7 6
great sundew (Drurmi m&u), carnivorous plmt 127 insularity 3 3 . 42
greater bird's-foot trefoil (/JXJIJ i t / i p i o ~ i i . ~helophyte
), 24 insularity. as r n v i r u n m r n t a l factor 33
green iguana (/piunu rprrnu). water loss 25 integrated pest management I36
grey hair-grass (Lotynephorrrr cunercenr), d i s t r i b u t i o n a n d interference (contest) c o m p e t i t i o n 114, 116, 136
temperature 55 i n t e r q x c i t i c c o m p e t i t i o n 11.4, 137
g r e y squirrel (S[irrriir [uru/inen.rir), in c o m p e t i t i o n w i t h red intraspecific c o m p e t i t i o n 114
squirrel 114 i n t r i n s i c rate of natural increase 86, 93
g r i z d y b r a r (Urrur on-tm),a n d disturbance 34-5; h o m e range inrrcxluced species 4 7 , 63. 77-82, I06
52-3 i r r u p t i o n s 87-8, 108
g r m s p r i m a r y p r d u c t i o n 143, 169 island biogeography, thcnry of 222
ground finches (Geuspizinae), character displacement i n 1 1 8 ; i v n r y gull (Pupphrlu eburned), coprophage I25
sanguinivory in 125
guild 16 jump dispersal 46, 47-8, 58, 7 0
j u n g l e m y n a (AL.rrdderrrrfNwr.r), and commensalism 1 1 .j
habitat islands 3 3
habitat: defined 13-15; diversity 161, 163; d i v e r s i t y hypothesis K - m i g r a t i u n strategists I 0 3
16.3; f r a g m e n t a t i o n 89, 97, 156, 169, 186, 208; generalists K-strategists 99-100, 1 0 1 , 108
15, 40; n i c h c 16; selection 5 3 , 5 5 ; specidists 15, 40 k a k a (Nertor nierrdion'h) /1, 7 . 12 I
halosere I74 kakapo (Strippf hu6roptif1(.1),avian browser a n d grazer 12 1
H a m i l t o n frog (kiope/iiru huinr/tonr), a n d i n t r o d u c e d predators 80 k a n g a r w rats (Dipodu?iry spp.), keystone herbivores 156; warer
h a r l e q u i n frog (Atelopirr wirirrr), a n d glubal w a r m i n g 198 i n t a k e 26
heat stress 21-2 kea (Nrrtur no/ubr/ri) 4. 7
heath cycle 174 keystone: herbivores 154; omnivores 15.4; predators 15.3. 15.1-6;
heath grass (Drintboniu de~-ririibear), st ress- tolerator 1 0 2 species 130, 153-8, 169
Hr/imrphur<rturri. carnivorous p i t c h e r - p l a n t I27 a n d commcns;ilism I 1 i
k i n g - c r o w (I1rz.rirrii.i ~ir'r<roceriiit),
heliotropism I 8 K i r t l a n d ' s warbler (Dendrorc.o krrr/'indii), thrratened b y g l o b a l
hclophytes 2 3-4 w a r m i n g 197
hemicryptophytes 17 koala (Ph'iiio/mfni mirrru~),arburral leaf-rarer I 56
herbivore niches 1 2 1
herbivore-plant systems 124 laclybird. LhrluiuriiJ n r p t i i l , accident.il dispersal 77
herbivory 120-5, 136 Lake M a n y a r a N a t i u n a l Park, n o r r h e r n Tanzania XX-!,
heterotrophs (consumers) 142, 145 land bridges 62
H i m a l a y a n tahr (Hrnrrrrqur j i ~ n r / u h i ~irruption ), in N e w land ethic 212
Z r a l a n d 87 l a n d igiranas ((.'nno/opbiir p ' h f i i r ) , overlapping liome ranges 5 2
historical biogeography 1, 5 , 63-4. 82 l a n d m a m m a l s ( i n Cenozoic era) 7 0
hoarzin (Opirthoiunrurhorrzis), avian browser 1 2 1 land-cover transformation 186
h o l l y (Ilex q i i r / o / r r i r i / ) , a n d l o w temperarure 5 5 landscape elements 13-14
h n m e range 52-3, 82, 98, I86 l a t i t u d i n a l d i v e r s i t y g r a d i e n t we diversity)
hmiensrasis 173, 223 l a w of t h e m a x i m u m 16
258 INDEX

law of the minimum I 5 mimicry: complexes 123; rings I I I

Leadheater's possum ( G ) i i i i i u h / i i l e i i i l e m / h e t i w i ) , in a fragmented mimics 1 11-12
landscape 9X mineral cycles 149
least shrew (Crjpto/iJ p m n i ) , individualistic response to climatic Mineral King Vallcy, Sierr.1 Nevacla. <:aliforni'i. wilrierness area
change 212
least weasel ( N i i i t r L i i i i i d ! ) , predator in Finland I 30 mineralization I 4 X , I69
climatic change 1x2, 18.4 mires, in Prince Edward 1sl;inds I99
Leitnerixeae, restricted endemic plant family -I.$ niwa ( D i o r i i i i iiidxiiiiiii), extinct large riirite 7 2
leopard (Puntber'i p r d i i i ) , Iwxlator i n southern India 128; i n mole riir ( ~ q p r o i i i yhomiitotii,), A w d avoider 2.1-5
Thiiland I 2 8 monoclim~x17 1-1
I.eiqwId, Aldo, conservationist 212 monriiiic klt 3 0
Liebig, Justus win, agricultural chemist I 5 mortalicy 84.XX. 01-1, 9X, 100
life table, 90, 91, I O X moth. Cliccubflritil L.xittiriiiii, biiiliigical <iinrrol agent I 3 5
life-fiirms 36-7. 'IL. I03 mountain pines (Piiiiir iiioiit,iii,i), i n Swiss Alps 30
limiting f ~ r i i r s15, 19, 2.3, 30, .it0 moiise ( A h viit.rtii/iti). prey swir~liing1 99
limiting resource(s) 11.1, 120. 128 Miillcrixi mimicry I 1 I
Ii t hobiomes 3 Z muntjiic deer ( ~ \ l i i i / t i u i i irtwteii),
~ in Britain 7X
litlimere 1 7 1 murualisni 109, I I ? . 136, 173
little bluesrem (.S&dqriiiiii i ~ ~ ~ i p ~ r i icompetitor
iiii), 120, 17X myrmecnpha~y(ant rating) 125
local popularinns X4, 95, 97, 1 3 3 , 1x9
logisric: eqmatiiin 8 7 ; grnwrh 86, 1 I 4 natality 8.1
liingleaf pine ( P i n i i i pufiiJ/rii), predicted distribution 56 h"itroiieb,a teriiiiti p q o i y i , alkal ipliile 3 0
loose metapol~iilatiiins94 n'itrerlack toad ( / j i i / r i ' i i l l r i i i i t l i )in Ireland 63; as i n c t : i p ~ i ~ ~ t i I a t i ~ i ~ i
Lntka, Alfred James. niideller of pupii1:ition dynamics I I.<, 1.30 97
Inwer critical temperature L O Kerophoiitri. relict Cariblxiii insectivore 7 i
lowland Atlantic fiiresr. Brazil 197. LIZ n c t primary p r ~ ~ l u ~ r iI.i'l-5. i ~ i i l.'I6, 1.47, 109
Lyme Park. Cheshire, England. red deer 1 q u l a r i n n 8.j-5 neutrnphiles iI
L p o w i i r i i J . Early Triassic mammal-like reptile 69-70 Newquay ZOO,Cornwall, England 21 I
nival zone 19, 3 0
m.icroconsumers (biophages) 146 no-modern-analogue cnmmunities IX2
macronutrients 147. 1-49 Noah's arks 62, 7 0
madder (Ritbui purqrin'i), northern limit 4-4 Northaw Great WcxxI. Hcrtfordshire, Ihgland. .is ecosystem
Magnoliacede, d climarically disjunct family -48 13x42
maidenhair spleenworr (Arpfmtiitii /ri~~hiiiii.iiiei), chasmophyte 3 1 northern fur seals ( C d l u r b i i i r r ~ itr.riiiiir),overexploited species
mainland-island metapopulmons 97 I04
male fern (D~optrri,r /i/ix-iii'r.i), stress-tolerant competitor I 0 2 northern plains pockcr gopher ( ' I ' b f ~ i ~ i ~t<i/piiicfuJ ~iiiji )-
malleefowl (Lripw mrNZitzr).in a fragmented landscape I 8 7 individu;ilistic response co c l i m a t i c change 182
Malthusian parameter 86 northern spotted o w l (Strix- o ~ ~ i d r t i ~ ,.mriii,i), drr in a fragmented
mammals, adaptations ro deserts 26 landscape 97
marine biomes 30 Niirway spruce (Piir'r ' i b r e i ) fnrest, in Swiss Alps 10
marine biosphere 2 8 Norwegian mugwort ( A r t e i r i i i i u iiiiri q i i ' i ) , climatic relict -4Y
marine iguanas (/\iiibljrhjiiJm iri~td/iir),population crash XX Noch?/;ipi (southern breches). disjunct group 7 I
miirram-grass ( Atiiiiinphil'i uri>iiliriti),pioneer I 7 -3
marsupials 4. X, 1 0 , 'I9, 6 5 , 77 omnivores 125. 151, 15.$-6, 1x5
mast fruiters 124 iirganochlnrines I 5 9
May, Robert bl.. discciverer of chaos in population dynamics oriental fruit moth (Gruphdith'i iiiulrrt~r).and integrared p s t
89 management I $6
mmclow oar-grass (/lulii/otrrthriii prurriiie), calcicole i I iiriibiomes 3 0 . 3 2 , 4 2
meadow pipit (Aiitbii~pr'iteiiiir), and habitat selection 55 oryx (Oiyx spp.). adaptations to heat Z I ; water intake 26
mesophyrcs 2 3 4 ) . cnmmensalism I I 3
otter (Liitnr i m ~ i d e i i ~ i iand
Afrwr'iiiriiJ, Permian reptile wtth disliincr distribution 69 otter (Liitr'i / i i / r d ) , threat frnm Ameriran mink in Britain 7 X
mesquite (Proiopi~spp.), ;idaptation to harsh substrate 3 I
metapopularinns 2. ij,8.t. 93-9. 108. 133. 1x6, 1x7: pale trefciil ('f'rifiliiiiii p~illrrieii.~).
protwcxyierator I 0 0
conservation 97-9 pandemics .13, X9
micrixonsumers (saprciphages) I46 Pangara 10. 38, 69-7.3, X 3
micronutrients 147, I49 pantropical distribution rue distribution
micro-endemics 4.5 paradox of enrichment I30
midge, Liputieiiru ~ i ~ i r r ~ i . r ~polysrenoxybionr
~riii, 16- I7 Pmwiiiruiitii, as protozoan prey species in laboratory experiments

migration strategies 103, I O X 86, 1 1 4 , 110. 132

milkweed (AJdepiuJmmrumd), piisonOtis plant 123 parasitism 19, 120
 INDEX 259
passenger pigeon (Ei~toprcter~n~grutorrr~r), hunted to extinction I04 rainbow covenant 217
passive replacement 76 rampant disperser 63
patch, definition of 14 range: 'creep' 63; contractian 63; expansion 63; size, explanation
pathogens 34, 35, 87, 100, 134 of 53; size and shape 51, 82
pedobiomes 32, 40 Rapqmrt's rule 52
peripheral species 197 ratites 48, 71, 72, 7 3
pest control 133-6, 158 Raunkiaer, Christen 37
phanerophytes 37 razorbill (Aolru tor&), auk survivor 104
photoperiod 17-18 realized (actual) niche 36
photosynthesis 16, 17, 11, 27, 100, 142, 145, 149 reciprocal co-occurrence pattern 80
Phyloloxeru, and the French wine industry 135-6 red ant, Aiynnri-u r u b u h , habitat requirements 15
phytomass 143, 146, 149 red deer (c'mwr ehphur): as population 84-6; hunting of 21 1
pikas (Oihotonu spp.), and global warming 197 red fox (Vrdpu tu/per): home range 52; predator of young
piiion lays (Gytnnorbrnur qunoiepbu/ur), seed predators 124 malleefowl 189
piiion pine pi nu^ eriulir), seed production i n 124 red squirrel (S~.rurrrrvdgurrr), competing with grey squirrel
plagioclimax Jer disclimax I14
plains pocket gopher (Geonryr burrurrrrr), humidity in burrow 26 red swamp crayfish (Procamburur ihzrkri), victim of
plant: associations 28; defences 121; formations 28, 30, 44, 173; biomagnification 159
life-forms 37, 38-9 red-osier dogwood (Cornur srooloni/ru), and acclimatization 22
pocket mice (Perognuthrrr), substrate preferences 3 1 reintroducing species 106
pikilotherms 19 relict insectivores 73-6
polar bear (Urrur nrurrtrnrr~r),adaptation to cold 20 religion, and Nature 218
Poluromonur w i - r d u t ~psychruphile
, 19 reptiles, adaptations to deserts 25
pliceman's helmet (Irnputrenr g/undrdr/eru), competitive ruderal resource partitioning 117, 136
102 reticulated velvet gecko (Oeduru rrtri-r~olrrtu), in a fragmented
polychlorinated biphenyls (PCBs) 159 landscape 189
plyclimax hypnrhesis 172 ring o u ~ (Turdur
l torquutur), climatic relict 4-5
Polynesian rat (Kuttur exuhnr), introduced predator 80 robin (Errthui~uurrubeculu me/ophi/rrr), survivorship curve 92
px)l frog (Rum lerronue), as metapopulation 96 rock hyrax: Heterohyrux brui-el, upper critical temperature 21;
population: age and sex structure 89; crashes 88-9; exploitation Heterohyrux and Procavru, saxicolous species 3 1
104-5; growth 84-92. 107, 108, 133; strategies 99, 100-3 rock wallabies (Perroguole and Petrodormr), saxicolous slxcies 3 1
pstclimax 171-2 Rtwky Mountain pika (Oihotonu prrncepr), saxicolous species 3 1
prairie wetlands, and global warming 198 Romanian hamster (Me~orrri~efrrr newton;), restricred endemic
preclimax 171-2 43
predator-prey cycles 129-33 rosebay willow-herb (Chumerron ungusrifooliunr). competitor 10 1
prey switching 128, 136, 199 rough-legged buzzards (Bufeo olugopiir), high immigration rate
prickly pear (Opiintra rtrti.fu), biolngical control of 134 88
primary sere (prisere) 174 ruderals 100-3
primary succession see ~uccession rust fungus (Pui~iniuubruptu var. purtheniicoolu),biological control
prisere ree primary sere agent 135
producers (autotrophs) 142
Proteaceae, grolugically disjunct plant family 48, 7 1 sabre-toothed: cats 156; tigers 65
protocooperation 109-1 1, 136; mutualism extreme form of sac fungus (Cryphonectrru (Endothm)purufrtrru). cause of chestnut
112 blight 8 1
Przewalski's horse (Equur przmdrkn), captive breeding saguaro cactus (Curnegreu gigantea), a succulent 27
programme 212 salt glands (in reptiles) 25
psammosere 174 saltarorial, adaptation in mammals 31. 37
psychrophiles 19 saprophages 142, 146, 147; fee U/JO decomposers
puma or cougar (Pelrr concoolor), pandemic 43 saprophytes 147
pyramid of numbers 152, 153 saprovores 142, 147
Pyrolobur fumurir, hyperrhermophile 19 saxicolous: animals 31; vegetation 30
scavengers 125, 145, 156
quackgrass (Agropyron repenr), good disperser 120 Sclater, Philip L. 10
quail (Coturnrx coturnix coturnrx), r-strategist 100 Sclater-Wallace scheme of biogeographical regions 1 0
queen scallop (Ch/umyr operru/urrr), and mutualism 112 scramble competition 114, 136
sea otter (Enhydra olurrrr), keystone predator 154
r-migration strategists 103 sea star (Prrurter ochrui-eur),keystone predator 154
r-strategists 99- 101 secondary production or zoomass (animal biomass) 146
radioactive isotopes, long-range transport of 158-9 secondary succession ree succession
rain shadow 33 secular migration 59
260 INDEX

seed p r e d a t i o n 124, 136, 1 5 6 tapirs .4-5. 66- 77

seres 173 tcchnocenrrics 2 1 3-15, 2 2 2
sharp-beaked ground f i n c h (G'eurptzd di/&ih~), sanguivnre technocentrism 213-1 5, 2 2 4
125 temperate (middle l a t i t u d e ) d i s t r i b u t i o n Ire distrtbution
sharp-shinned h a w k (All-ipitrrrtrrutilr), a n d resource p a r t i t i o n i n g t e r r i t o r y 53. 63. 97* 98, 1 1 4
I17 thalassochore 59
shelf f u n g u s (T'rutirefeJwrJico/ur), decomposer 147 t h e r m a l n e u t r a l zone 20, 56
shepherd's p i i r s e (C'd/IJd/ublL~.rU-,fIUJtorrJ), ruderal 102 t h e r m o p h i l e s 17. 19
Sierra Club 2 1 2 therophytes 25, 17,1 7 8
silverweed (Purrntr//u mrerrm), a n d geese 1 2 5 t h r e e - t d w i x d p e c k e r (Piloidrr frrdu'fy/irJ),cool t r n q x r a r e forest
s k i p p e r b u t t e r f l y (HeJperru W U I M ) , in f r a g m e n t e d l a n d s c a p dweller 15
IX 6 t h y l a c i n e o r Tasmanian w o l f , e x a m p l e of cimvcrgenr e v o l u t i o n
s l o p e (inclination), as e n v i r o n m e n t a l factor jj 65
small-leaved l i m e (Tilru mrduiu): w a r m t h r e q u i r e m e n t 22; ticklegrass or hairgrass ( A p J t i . rnabru): p o p u l a t i o n crash 89;
n o r t h e r n limit 5 5 gold disperser I20
snow 27, 30 t i g c r (Puntheru ti,qrij), predatur in southern I n d i a I 2 X
snowshoe hare (Lepm t i l t ? u ~ / e ~ ten-year ~ ~ ~ ~ r ) ,p o p u l a t i o n cycle 129 tight m e t a p o p u l a t i o n s 94-5
Snay sheep ( 1 h urrer). p o p u l a t i o n crash X8 tolerance m o d e l (or succession) 17 3 . 208
aocial I > ; i n v m s m 21 5, 223 t o l r r i i n c e range 16, 4 0
solar r a d i a t i o n 17, 184 t r a p p i n g 77, 107. I O X
Sofrviodun, r e l i c t Caribbean i ~ i ~ e c t i v 7~ ir e tree p i p i t (Atrthii~/ r r i d r ! ) ,a n d h a b i t a t selection 55
Spaceship E a r t h 221, 2 2 2 tree-r h r o w 3 5
spatial: compc.ritiun hypnthesis 120; m o d e l 193 ruatara (S/~hetror/oa/Irmc.iufii.r),r e l i c t NKW Zealand r e p t i l e 49,
spccies, defined 6; c(xxistencc 116; d i v e r s i t y 161, 163-6; u n d e r XO
pressure 197
species-area: curve 161, 169; effect 161; relationships 161 u m b r e l l a thorn ( A m r u turti/iJ), a n d disturbance 89
speckled rangeland grasshopper (Arphi'r L . o ~ s ~ ~ d~i sJ tur i)b,u t i o n u p p e r c r i t i c a l temperature 2 I
and habitat requirements 4
. S ~ ~ U ~ N(I hI g I moss) 33, 45, 110, 172, 176-7 vagiliry 5
spinifex hopping mouse (Nutony ulexrr), w o r l d c h a m p i o n u r i n e vegetatiun ri)posequences 33
concentratnr 2 6 vegetation succession J C succession ~
spruce ( P m u ) , r - m i g r a t i o n strategist 103 v e r n a l i z a t i o n 22
s t a n d i n g c r n p 143 vicariance: 2, 5. 8, 10, 60, 67, 73, 8.5;biogeography 5, 67;
stemless t h i s t l e (C'rrJiimi ucuide), n o r t h e r n limit 55 events 8, 10. 60, 7 3
s t e p p i n g stone islands 62, 70, 98, I ? ? vicars 36, .42, 77
s t r a t i f i c a t i o n (of c o m m u n i t i e s ) 141 Victorian Acclimatization Acclimatizarron Soorty. Ausrralia
strawberry tree (ArhirtrrS irnedo), long-day p l a n t I 8 210
stress, defined 16 Viking funeral ship 76
stress-tolerant c o m p e t i t o r s 1 0 2 vnles (Alil-ruti~~ spp.), prey species in F i n l a n d 1 3 0
stress-tolerant ruderals 1 0 3 Volterra, Vito, p o p u l a t i o n m n d r l l e r 114. 130
stress-tolerators 39, 100-3, 108
subalpine lxlt 30 w a l l f l o w e r (C'hrirutrthrrr cberri), c h o m o p h y t e 3 I
s u b c l i m a x 171 Wallace, A l f r e d Kussel. biogeogrdpher 10. 63
submontane belt 30 w a n d e r i n g albatross (Dromedid exu/utr.t),94, 100
substrate, as e n v i r o n m e n t a l factor 30-2 warblers (genus Detrdruicu), a n d resource p a r t i r l o n i n g I 17
succession: 17 1-3; allogenic 174; autogenic 174; in abandoned water stress 21, 201
fields, M i n n e s o t a 178; in G l a c i e r Bay, Alaska 174, 181; in water tupelo ( N ~ J Juqirutr~.u), U w e t tolerator 24. 191
K r a k a t a u Islands, Indonesia 177, 181-2; in western G r e a t water v o l e (Anwo/u termtrir), d e c l i n e in B r i t a i n 7 8
Basin g h o s t t o w n s 178; m u l t i d i r e c t i o n a l 181-2; p r i m a r y 174, w a v y hair-grdss ( ~ r J ~ ~ h d f u/ & / Ii~~ ,ijud ) , cdtclfilgr 3 I
181, 208; secondary 174, 178-80, 2 0 8 wetland. loss of 189
succulents 25, 27, 36, 37 w h e a t r d r (Octiunrhe uenutrfhe), a n d h a b i t a t selection 5 5
Siilfhbii~uciduc'i/durii/J,h y p et t h e r m o phi I e I9 w h i t e a d m i r a l b u t t e r f l y (Ludoxu ~ i r m r / / u ) response
, to c l i m a t i c
s u l x r t r a m p s 60-1 w a r m i n g 197
SulVlVdl rates 90-4 w h i t e f l y , Poruben/tJiu t q r i w e , c i t r u s pest 135
s u r v i v o r s h i p curves 91-2, I O X Wild F r e e - R o a m i n g H o r s e a n d Burro A c t ( 1 9 7 1 ) 21 1
s u s t a i n a b i l i t y 222-3 w i l d l i f e trade 167, 169, 21 I , 2 2 4
sweepstakes routes 6 2 wooclrats (Neimmu spp.), adaptation to remixxrarure 2 1
Swiss stone p i n e s (PIIWJmrbru), in Swiss A l p s 3 0 W y r h a m W o o d , Oxfordshire, E n g l a n d . fuod w e b i n 151

t i i n n i n s (in oaks) 1 2 3 xerophytes 23, 25, 27

 INDEX 261

xerophytic adaptations 25 Yorkshire fog (HO/UJInnutuj), C-S-R strategist 103

yapok or water opossum (Chronecter ~ ~ Z ~ Z ~ Z Z aquatic

LJ), marsupial zebra finch (Tueniopygru rsrrtunott~),av~anr-strategist
66 100
yellow rajah-rat (Muxonzyr uri,&), prey species in southern India zomass 146
128 zonobiornes 27-8, 30, 32, 4 0
yellow-bellied marmot (Mummufirswntns), saxicolous species zonoecotones 30
31 z w s 211-12. 224