CARNIVOROUS PLANTS

of the United States and Canada

by Donald E. Schnell
CARNIVOROUS PLANTS
of the United States and Canada
Sarracenia flava

JOHN F. BLAIR, Publisher

Winston-Salem, North Carolina

CARNIVOROUS PLANTS
of the United States and Canada

by Donald E. Schnell
Copyright © 1976 by Donald E. Schnell
Library of Congress Catalog
Card Number: 7 6 - 2 6 8 8 3
All rights reserved
ISBN 0 - 9 1 0 2 4 4 - 9 0 - 1
Printed in the United States of America
by Lebanon Valley Offset Company Incorporated

Library of Congress Cataloging in Publication Data

Schnell, Donald E 1936-

Carnivorous plants of the United States and
Canada.

Bibliography: p.
Includes index.
1. Insectivorous plants—United States. 2. I
sectivorous plants—Canada. I. Title.
QK917.S36 583'.121'0973 76-26883
ISBN 0 - 9 1 0 2 4 4 - 9 0 - 1
Sarracenia psittacina
For Lee Anne
& Kristen
Foreword
By C. RITCHIE BELL
Professor, Department of Botany,
University of North Carolina, Chapel Hill
and Director, North Carolina Botanical
Garden, Chapel Hill

Since their discovery by b o t a n i s t s over two hundred of the plant world f r o m rapid extinction through a
years a g o , the world's carnivorous plants have b e e n the c o m b i n a t i o n of realistic protection f r o m continued col-
center, f r o m time to t i m e , of m u c h scientific study and lecting, conservation of their specialized habitats,
public interest. T h e y have also b e e n the center of a responsible horticultural research, and the general cul-
n u m b e r of m i s c o n c e p t i o n s a n d , in the past few y e a r s , tivation of only those plants actually grown f r o m seeds
of considerable commercial exploitation. o r , w h e r e p o s s i b l e , cuttings.
C a r n i v o r o u s plants are b o t h colorful and biologi-
C. R . B .
cally interesting, but they are not h o u s e plants. T h e i r
July, 1976
specific e n v i r o n m e n t a l requirements o f t e n involve very
high humidity, high light intensities, quite acid soil,
pure w a t e r , and seasonal temperature c h a n g e s ; such
requirements are not met by a pot of generalized dirt,
treated tap w a t e r , and the usual temperature and h u -
midity ranges of the average h o m e , office, or apart-
m e n t ! It is likely that not o n e " c u l t i v a t e d " carnivorous
plant in a thousand lives a full year and p r o b a b l y less
than one in ten t h o u s a n d survives f o r two years in the
hands o f most a m a t e u r " c o l l e c t o r s " o r those w h o , u n -
f o r t u n a t e l y , are misled by the c o m m e r c i a l advertise-
m e n t s f o r these unique plants and attempt to g r o w
t h e m , merely as a n o v e l t y , on the k i t c h e n w i n d o w sill.
G i v e n the a b o v e b a c k g r o u n d , this b o o k b e c o m e s
s o m e t h i n g of a l a n d m a r k in carnivorous plant p u b l i c a -
tion. N o t only does it bring together in a very readable
f a s h i o n the m a n y interesting b o t a n i c a l facts c o n c e r n i n g
the f o r m , f u n c t i o n , and ecology of our carnivorous
plants, but the a u t h o r m a k e s a s t r o n g , and very timely,
case for the p r e s e r v a t i o n of these unique m e m b e r s

vi / Carnivorous Plants of the United States and Canada

Preface
T h i s b o o k is intended f o r practical use. It is not c o n -
ceived of as a coffee table or b o o k c a s e o r n a m e n t ; the
entire plan and structure of the w o r k is centered around
utility.
T h e p h o t o g r a p h s have b e e n c h o s e n with a view
toward sharpness of important f e a t u r e s , clarity, color
fidelity, and m o s t of all, h o w well they represent the
plant. Y o u will find only a few drawings in this b o o k ,
and these are mainly of a figurative nature. T h e trouble
with using only botanical drawings f o r identification is
that they represent a t w o - d i m e n s i o n a l , colorless a v -
erage of characteristics in an ideal plant that rarely
exists. T h i s presents a p r o b l e m f o r the b e g i n n e r in the
field, who has a b o o k of drawings in h a n d but sees no
plants that resemble any of the d r a w i n g s , or perhaps
sees too m a n y .
T h e p h o t o s are intended to show the plants as they
really are, and m o s t of the pictures have b e e n made in
the field. If some of the p h o t o g r a p h s seem p r e t t y or
artistic, that is just a b o n u s . S i n c e no view of a plant
or group of plants can show all the important c h a r a c t e r -
istics one might wish to see, I have presented multiple
views of them where n e c e s s a r y .
O c c a s i o n a l l y , in the field, where the natural b a c k -
grounds tended to camouflage, I h a v e had to resort to
the use of a neutral gray b a c k g r o u n d , reflectors, or a n -
cillary lighting to m a k e the s u b j e c t s stand out properly.
O n l y a few pictures were made under studio condi-
tions, using live plants f r o m my collection. A f t e r h a v -
ing observed, g r o w n , and w o r k e d with carnivorous
plants f o r eighteen y e a r s , I felt capable of m a k i n g a
proper selection of cultivated plants for p h o t o g r a p h y .
I have departed f r o m the traditional botanical s c h e m e
of presenting species within a genus in alphabetical or
other order, preferring to start with some c o m m o n ,
vii / Preface
m o r e readily o b s e r v e d plants or those that would b e s t
illustrate general characteristics, or p e r h a p s grouping
similar species together so that c o m p a r i s o n s could be
seen.
T h e carnivorous f u n g i have b e e n excluded. T h e b o o k
deals only w i t h the green seedplants. Fungi require c u l -
ture and m i c r o s c o p e s f o r proper o b s e r v a t i o n , and the
species are in a state of t a x o n o m i c disarray at present.
W h i l e f a s c i n a t i n g and certainly deserving of f u r t h e r
s t u d y , t h e y are not ready f o r this sort of presentation.
T h e text is not c o m p l e t e in the classical botanical
sense of a m o n o g r a p h , but it w a s not intended to be.
O n the o t h e r h a n d , the occasional o r b e g i n n i n g n a t u -
ralist m a y find m o r e t h a n he needs and can easily be
selective in w h a t he reads. T h o s e interested in f u r t h e r
study will find r e f e r e n c e s on m a n y levels at the end
of the b o o k .
• • •
I would like to t h a n k J o h n F. Blair, Publisher, and
his staff f o r taking on this project in the first place,
and then seeing it t h r o u g h to a quality production.
W a r r e n P. S t o u t a m i r e was kind e n o u g h to read p o r -
tions of the text and offer m a n y useful s u g g e s t i o n s .
Letters and discussions w i t h Peter T a y l o r and K a t s u -
hiko K o n d o were very helpful to me in developing my
tentative concepts of the difficult t a x o n o m y of Utricu-
laria, b u t I b e a r full responsibility f o r the system
presented in C h a p t e r 7, including errors. Finally, I
offer tenderest appreciation to my w i f e B r e n d a , w h o
has b e e n patient and encouraging during the p r e p -
aration o f this b o o k and w h o has a c c o m p a n i e d m e
on m a n y field trips, o f t e n serving as my stern e a r l y -
warning system f o r potentially dangerous reptiles,
q u i c k s a n d , and treacherous b o g h o l e s .
Contents
Foreword vi Drosera anglica 61
Drosera intermedia 63
Preface vii Drosera filiformis 64
Drosera capillaris 67
Chapters Drosera brevifolia 69
Difficult Identifications 70
I. Carnivorous P l a n t s — A n Introduction 1
Carnivorous or Insectivorous P l a n t s ? 4 V I . T h e B u t t e r w o r t s (Pinguicula) 71
Kinds of T r a p s 4 Pinguicula vulgaris 74
Carnivorous Plants around the C o n t i n e n t 6 Pinguicula pumila 76
T h e Future of C a r n i v o r o u s Plants 9 Pinguicula lutea 77
Flower Structure 10 Pinguicula caerulea 78
How Plants Are N a m e d 13 Pinguicula planifolia 78
Pinguicula primuliflora 79
II. T h e V e n u s ' Flytrap (Dionaea muscipula) 16 Pinguicula ionantha 80
Pinguicula villosa 81
III. T h e Eastern N o r t h A m e r i c a n Difficult Identifications 82
Pitcher Plants (Sarracenia) 22
Sarracenia purpurea 28 V I I . T h e Bladderworts (Utricularia) 83
Sarracenia flava 31
VIII. Growing North American
Sarracenia alata 34
C a r n i v o r o u s Plants 96
Sarracenia oreophila 36
G e n e r a l Principles 96
Sarracenia minor 37
Genus Notes 106
Sarracenia psittacina 40
An Outdoor Home Bog 109
Sarracenia leucophylla 42
S o m e C o m m e r c i a l Sources o f
Sarracenia rubra 44
C a r n i v o r o u s Plants 111
Sarracenia Hybrids 47
Field Collecting 112
Difficult Identifications 51

Additional Reading 114

IV. T h e California Pitcher Plant
(Darlingtonia californica) 52 Glossary 118

V. T h e S u n d e w s (Drosera) 56 Derivations of Scientific Names 121

Drosera rotundifolia 58
Drosera linearis 59 Index 124

ix / Contents
I. Carnivorous Plants—An Introduction
T h e concept of a plant that traps and partially c o n -
sumes small animals was suggested and studied long
before 1 8 7 5 , w h e n D a r w i n ' s b o o k Insectivorous Plants
appeared. In that volume D a r w i n correlated a great
deal of the k n o w l e d g e of his day and added the results
of his own n u m e r o u s experiments and o b s e r v a t i o n s .
Since t h e n , interest in carnivorous plants has g r o w n
r e m a r k a b l y . C o n c u r r e n t l y with this g r o w t h , new dis-
coveries and ideas have arisen, and these have further
increased interest in carnivorous plants. Imaginations
have also soared, but stories of giant or m a n - e a t i n g
plants have proved to be entirely mythical.
G r e e n plants can derive a large part of their chemical
requirements for sustaining g r o w t h and reproduction
from some very basic but essential e l e m e n t s . T h e s e
include energy f r o m sunlight, o x y g e n and c a r b o n di-
oxide f r o m the air, and water and certain minerals
from the earth or water in which the plants grow.
T h r o u g h p h o t o s y n t h e s i s , in which green chlorophyl
plays so large a part by t r a n s f o r m i n g sunlight into
chemical e n e r g y , c a r b o h y d r a t e s are built up f r o m water
and carbon dioxide. T h e s e various c a r b o h y d r a t e s t h e m -
selves are used as energy sources and building b l o c k s
for synthesizing myriad other chemical materials
needed by the p l a n t — p r o d u c t s such as amino acids
and proteins, vitamins and h o r m o n e s , and even small
quantities of fats.
During the entire complex process, which may be
likened to an automated chemical f a c t o r y that goes on
and on as long as raw materials are supplied and end
products r e m o v e d , m a n y minerals are a b s o r b e d by the
plant's root system for ultimate inclusion in chemical
end products. Examples of such minerals are c o m -
pounds of nitrogen, p h o s p h o r u s , p o t a s s i u m , calcium,
1 / An Introduction
m a g n e s i u m , iron, m a n g a n e s e , b o r o n , and several other
elements needed in such m i n u t e quantities that they
are seldom deficient in the e n v i r o n m e n t .
D u r i n g millions of years of evolution, plants h a v e
s h o w n a clear capacity for adapting to different h a b i -
tats which may be deficient in one or more of the basic
requirements. For e x a m p l e , certain plants are able to
grow and function very well in the deep shade of m a -
ture f o r e s t s , where sunlight is quite reduced, because
their p h o t o s y n t h e t i c apparatus f u n c t i o n s more effi-
ciently than that of o t h e r plants. N o t only do such
shade-loving plants thrive on the forest floor, but they
often do poorly if exposed to full sunlight. As another
e x a m p l e , in dry desert regions all over the world plants
have adapted to very low water levels t h r o u g h o u t
much of the year. T h i s is accomplished by various de-
v e l o p m e n t s : deeper root s y s t e m s , chemical monitoring
of the spacing of plants by which no one plant a b s o r b s
too m u c h water f r o m a unit of earth, reduced rates of
water loss by c h a n g e s in leaf c h e m i s t r y and s h a p e , or
the adoption of a life cycle that fits into the brief period
of annual d o w n p o u r w h e n the desert b l o o m s and
plants are able to g r o w and reproduce very rapidly
b e f o r e the land dries out again. T h e r e are m a n y more
examples of adaptation to w h a t for m o s t plants might
be termed difficult e n v i r o n m e n t s .
T h e kinds of deficient h a b i t a t s that will mainly c o n -
cern us t h r o u g h o u t this b o o k are the usually acid,
m i n e r a l - p o o r b o g s and s w a m p s , and the f r e s h w a t e r
m a r s h e s and s a v a n n a h s . It is in such locations that
plants that h a v e adopted carnivory m a y be found.
A n y o n e w h o sees a f r e s h w a t e r b o g or s w a m p is i m -
pressed by the great variety of plant l i f e — i n c l u d i n g
m a n y m o s s e s , f e r n s , and o r c h i d s — a n d b y the s e e m -
ingly rich, black ooze of the wet soil in w h i c h the plants
g r o w . But accurate chemical analysis of the soil in such
areas f r e q u e n t l y reveals that this initial impression is
partly e r r o n e o u s . First of all, the c o f f e e - b r o w n waters
are very acid, and acid w a t e r along w i t h f r e q u e n t
drenching rains quite efficiently leaches out m a n y ir-
replaceable minerals. S e c o n d , in w a r m climates there
is a very high rate of bacterial and o t h e r m i c r o o r g a n i c
activity w h i c h in itself uses up the sparse supply of
minerals that are needed by the larger plants. In cool
climates such decay is delayed, b u t then the undecayed
remains of dead plants and animals keep the m u c h -
needed minerals locked up. T h i r d , a close e x a m i n a t i o n
in m o s t cases confirms that the b l a c k ooze is little more
t h a n fine white sand along with a great deal of c h e m i -
cally sterile c a r b o n or c h a r c o a l - l i k e m a t e r i a l , or the
latter without sand. It seems that in such m i n e r a l - p o o r
h a b i t a t s there m u s t h a v e b e e n considerable adaptation
by all the plants in order f o r t h e m to g r o w and r e -
produce.
To adapt to these e n v i r o n m e n t s deficient in m a n y
minerals and possibly in some cases to o v e r c o m e the
inability of their root s y s t e m s to a b s o r b required m i n -
erals, some plants retained the evolved capacity to
trap and digest small animals. From the trapped ani-
m a l s , which are largely i n s e c t s , the plants a b s o r b w h a t
t h e y need. As a result of selective a d a p t a t i o n , the plants
that were able to trap animals survived and produced
offspring with the same genetic characteristics.
T h e acceptable w o r d " p r e y " is usually used in r e f -
erence to the entrapped a n i m a l s , b u t it is n o t literally
correct in that the plants do n o t actively stalk and c a p -
ture food in the sense that m a n y animals do. R a t h e r ,
the plant is able to lure or take a d v a n t a g e of incidental
nourishing visitors b y m e a n s o f one o f four k i n d s o f
trapping m e c h a n i s m s , w h i c h will be discussed b e l o w .
2 / Carnivorous Plants of the United States and Canada
After e n t r a p m e n t , the prey undergoes digestion.
F r o m a chemical standpoint the digestive process is
quite similar in m a n y respects to digestion in animals.
A l s o , various m i c r o o r g a n i s m s such as bacteria aid
m a n y carnivorous plant species in b r e a k i n g down the
prey into simpler, a b s o r b a b l e substances.
In the decades immediately b e f o r e and after the turn
of the c e n t u r y , m a n y e x p e r i m e n t s were contrived to
prove that digestive activity actually occurs in plant
traps and to m e a s u r e and define the nature of that a c -
tivity. S o m e of these e x p e r i m e n t s were quite elaborate
and their results still stand. For digestion to occur,
certain e n z y m e s must be present. E n z y m e s participate
in the chemical reactions of biological organisms by
causing the reactions to be completed rapidly at t e m -
peratures suitable for the m a i n t e n a n c e of life. T h e s e
reactions include the synthesis of more complex c o m -
pounds as well as the reduction that occurs in diges-
tion. T h e results of m a n y experiments indicate that
e n z y m e s are responsible for digestion in the traps of
carnivorous plants.
T h e n e x t question concerns the source or sources of
these e n z y m e s : D i d they originate f r o m the plants
t h e m s e l v e s , secreted into the trap along with fluid as
a response to entrapped prey of a suitable n a t u r e ; or
were t h e y simply products of bacteria or fungi i n h a b -
iting the decaying detritus accumulated in some open
t r a p s ? Like m o s t questions of this nature in science,
a categorical " e i t h e r - o r " a n s w e r is impossible, and it
would be misleading to attempt to give one. It has been
s h o w n that some species of carnivorous plants have a
complex e n z y m e - s e c r e t i n g s y s t e m in small, specialized
plant glands associated with the trap. O t h e r s with
similar glands secrete practically no enzyme under
sterile experimental conditions where the contribution
o f any m i c r o o r g a n i s m can b e discounted. A n d some
plant traps function with no glands at all. At this stage,
the answers are far f r o m complete. S o m e plants seem
to rely almost exclusively on their own e n z y m e s , some
seem to depend almost totally on bacterial action, and
others take advantage of b o t h sources.
A n o t h e r question concerns w h a t digestive products
are actually a b s o r b e d by the trap of the carnivorous
plant, which of these are truly required by the plant,
and which are just passively absorbed. A second, r e -
lated question is whether all the useful materials a b -
sorbed by the plant are simple minerals which m a y be
lacking in the plant's h a b i t a t , or w h e t h e r some are
more complex, synthesized materials needed b e c a u s e ,
as a result of evolutionary c h a n g e , the plant has lost
the capacity to produce t h e m . T h e individually studied
cases are few and far f r o m c o m p l e t e , so again we can
give only some partial a n s w e r s — m e r e l y clues in a
highly complex problem that involves more than curi-
osity about carnivorous plants and actually cuts across
the whole problem of the nature of adaptation.
Of all the mineral elements m e n t i o n e d previously,
the one that green plants need most consistently and
in the largest amount is n i t r o g e n , followed by p h o s -
phorus and potassium in m o r e variable quantities. Acid
soils are also quite deficient in calcium. All these ele-
ments are retained by " s w e e t " or basic s o i l s — t h u s the
gardener adds lime (a calcium c o m p o u n d ) to " s w e e t e n "
or enrich soil that is too acid to permit m o s t plants to
grow well. M u c h research has centered on the idea that
nitrogen is the limiting f a c t o r , or element m o s t needed
by carnivorous plants for sustenance and g r o w t h in
their deficient e n v i r o n m e n t , p r o b a b l y because nitrogen
has long been prominent in soil and plant c h e m i s t r y .
But insufficient w o r k has b e e n done to establish the
exact role that some other minerals or c o m b i n a t i o n s of
minerals may play in plant carnivory. For e x a m p l e ,
3/ An Introduction
recent preliminary o b s e r v a t i o n s indicate that p o t a s -
sium levels in soils, plants, and prey greatly influence
the a m o u n t and rate of nitrogen a b s o r p t i o n by c a r n i v -
orous plants.
V e r y early a t t e m p t s were made to find out w h e t h e r
a n y t h i n g could actually be a b s o r b e d by carnivorous
plants. R e s e a r c h e r s utilized h a r m l e s s dyes which could
be followed visually in their course through the plants.
T h e air s u r f a c e s of m o s t plants are covered by a thick,
w a x y layer called cuticle. T h e absorption of w a t e r y
materials t h r o u g h a w a x y layer of cuticle varies f r o m
slow to impossible. It w a s noted quite early that the
a b s o r b e n t interior s u r f a c e s of the traps of carnivorous
plants lack cuticle. It t h e r e f o r e was possible to follow
the dyes visually in their course t h r o u g h the plants.
T h e s e were i m p o r t a n t preliminary results. O f course
the e x p e r i m e n t s had very severe limitations.
Later, with the advent of radioisotope tracers
wherein various portions of a material can be tagged
with radioactivity and followed through the plant and
in actual chemical c h a n g e s in plant tissue, it was pos-
sible to conclude that absorption of certain materials
did take place and that these materials were actually
used by the plant t i s s u e s — t h a t is, the s u b s t a n c e s did
not just passively enter the plant tissues. So far these
studies h a v e been limited to n i t r o g e n c o m p o u n d s , and
we have only the published reports of studies by one
w o r k e r using one species of carnivorous plants out
of the forty or so on this c o n t i n e n t a l o n e ; but it is a
b e g i n n i n g , and it is certainly indicative that carnivory
must be of some benefit to the plant.
Additional w o r k on m o r e general levels suggests
that some carnivorous plants can subsist without
trapped and digested a n i m a l s , or that m i n u t e q u a n t i -
ties of suitable fertilizers can be substituted by apply-
ing them to the r o o t s , the trap interiors, or even the
external leaf surfaces. H o w e v e r , a c o m m o n o b s e r v a t i o n
in such e x p e r i m e n t s is that t h e p l a n t s are n o t as
vigorous as in n a t u r e : t h e y g r o w m o r e slowly and do
not b e c o m e as l a r g e ; they are m o r e p r o n e to d i s e a s e ;
and very i m p o r t a n t l y , they do n o t reproduce as well,
as is indicated by the production of f e w e r flowers and
seeds, a reduced rate of seed m a t u r a t i o n , and less
rhizome budding.
S o f a r , w e h a v e looked a t c a r n i v o r y f r o m the v i e w -
point of an isolated, e x p e r i m e n t a l plant. But plants oc-
cur in nature with o t h e r similar and dissimilar p l a n t s ,
with a n i m a l s , and with an i n a n i m a t e e n v i r o n m e n t as
parts of a c o m m u n i t y . T h e r e results a complex i n t e r -
action of so m a n y f a c t o r s that one is awed and baffled
in one's first a t t e m p t s to picture the situation in p e r -
spective. T h e picture is f u r t h e r complicated by the f a c t
that biological c o m m u n i t i e s are not s t a t i c ; they are al-
w a y s v a r y i n g and responding to assault and c h a n g e .
W h e n a prime e n v i r o n m e n t f o r c a r n i v o r o u s plants
c h a n g e s f r o m wetland to g r a s s y field, s c r u b , or forest
as a result of natural or m a n - m a d e activities, c a r n i v -
orous plants and m a n y of their w e t l a n d c o m p a n i o n
species disappear s o m e w h a t p r o m p t l y , o f t e n in a rather
specific order. T h e y are a p p a r e n t l y crowded out by
f o r m s more vigorous and b e t t e r adapted to w h a t has
b e c o m e essentially a n e w e n v i r o n m e n t . It seems that
carnivorous p l a n t s require the poor soils of an acid
wetland to be c o m p e t i t i v e , soils w h e r e m a n y other
plants that under different conditions would be strong
competitors c a n n o t g r o w . W h e n dryland plants that
d e m a n d richer soils are finally able to spread into a r e -
duced b o g or m a r s h , c a r n i v o r o u s plants b e c o m e the
disadvantaged f o r m s and disappear.
T h i s is n o t so difficult to u n d e r s t a n d or accept in
b r o a d t e r m s if v i e w e d f r o m a simplified b u t largely
valid e v o l u t i o n a r y angle. W e b e g a n this section b y
m e n t i o n i n g the adaptation of plants to differing sit-
4 / Carnivorous Plants of the United States and Canada
u a t i o n s . N o t all plants able to g r o w in the environment
of an acid, mineral-deficient wetland adopted carniv-
ory. Evolution seldom n a r r o w s to one p a t h w a y or one
structural adaptation to solve a problem. Variation and
gradual m i g r a t i o n are the k e y s to the continuation of
some life f o r m s in a continually changing environment.
CARNIVOROUS OR INSECTIVOROUS PLANTS?
I will not b e l a b o r the point as to w h e t h e r these plants
should be called " c a r n i v o r o u s p l a n t s " or " i n s e c t i v o r o u s
p l a n t s , " but I will m e n t i o n it lest the reader b e c o m e
c o n f u s e d by the use of b o t h terms in conversation or
in o t h e r publications. W h e n carnivorous plants were
first noticed and studied, the m o s t obvious prey was
insects; hence the term insectivorous plants. Later,
species with m o r e varied appetites were found. S k e l e -
tons of small birds and amphibians were found in some
traps, and aquatic plants trapped small water animals
that were clearly not insects. T h u s the term carnivorous
plants w a s coined to be more general and inclusive,
and m o r e accurate. It is the preferred term and the one
we shall use t h r o u g h o u t this b o o k .
KINDS OF TRAPS
T h e traps of carnivorous plants are modified leaves
that in some cases are so changed and adapted to their
f u n c t i o n that t h e y resemble only remotely leaves as
m o s t people picture t h e m . For e x a m p l e , the tall, often
decorative tubular pitcher leaves of species of Sarra-
cenia are f r e q u e n t l y t h o u g h t by the uninitiated to be
flowers, and in m o s t cases the trap leaves are far more
striking than the p l a n t s ' true flowers. T h i s case of
m i s t a k e n identity is s o m e w h a t ironic since it is a c -
cepted that m o s t true flowers are decoratively struc-
tured in order to attract insects or other animals as
pollinators. T h e trap leaves are also attractive to i n -
sects, b u t f o r a different end.
 T h e r e are four types of traps in seed-bearing c a r n i v -
orous plants of our region, and I have further divided
these into two main g r o u p s , active and passive. I would
reiterate that " a c t i v e " is used in a restricted s e n s e , not
as it might be used in c o n n e c t i o n with animals of prey.
A classification of these trap f o r m s along with e x -
amples is in outline f o r m below and can be correlated
with the a c c o m p a n y i n g p h o t o g r a p h s and d r a w i n g s .
Active t r a p s . — T h o s e in which s o m e rapid plant m o v e -
ment takes place as an integral part of the trapping
process.
1. Closing t r a p s . — T h e s e are o f t e n referred to e r r o -
neously as of the beartrap type. T h e trap is b i v a l v e d ;
that is, it has two similar halves c o n n e c t e d by a m i d -
rib. T h e two halves close on e a c h other and thus trap
the prey. T h i s type is represented in the w e s t e r n h e m -
isphere by only one species, Dionaea muscipula (the
V e n u s ' flytrap).
2. T r a p d o o r s . — T h e s e are aquatic traps, relatively
minute, and are represented by the g e n u s Utricularia
(the bladderworts). T h e trap is s o m e w h a t b u l b o u s ,
with a flaplike door over a small e n t r a n c e at o n e end.
T h e stimulation of sensitive external trigger hairs near
the trap entrance results in the opening of the door and
an inrush of water with the prey. A f t e r w a r d s , the door
closes again.
Passive t r a p s . — T h o s e in w h i c h rapid plant m o v e m e n t
is not an integral part of the trapping process.
3. P i t f a l l s . — T h e s e are characteristic of the familiar
pitcher plants of the genera Sarracenia and Darling-
tonia. T h e leaves are tubular with various o t h e r m o d i -

Fig. 1-1. Dionaea muscipula, the Venus' flytrap, with a

trap of the closing type.

Fig. 1-2. Utricularia gibba, a bladderwort, with a trap of

the trapdoor type. The plant is aquatic and the bulblike
trap is only 2-3 mm.

5 / An Introduction
 fications. T h e p r e y is lured to the pitcher opening,
enters or falls in, is u n a b l e to escape, and is digested.
4. " F l y p a p e r " or adhesive traps.—These occur in
Drosera ( s u n d e w s ) , and Pinguicula (butterworts). Nu-
m e r o u s sticky glands cover the upper leaf surfaces,
and the small prey is immobilized by b e c o m i n g mired
down. A f t e r e n t r a p m e n t , the stalked glands of Drosera
do o f t e n m o v e slowly and there frequently is some
slow leaf folding in some species, but this is part of
the digestive r a t h e r than the e n t r a p m e n t process.
T h i s brief outline is f o r o r i e n t a t i o n ; details of vari-
ous traps and their activities will be discussed in the
ensuing chapters.

CARNIVOROUS PLANTS
AROUND THE CONTINENT
W e have noted that carnivorous plants occur main-
ly in acid, f r e s h w a t e r w e t l a n d s . As is the case with all
g e n e r a l i z a t i o n s , this o n e h a s an exception or two.
Drosophyllum luscitanicum, a native of Portugal and
parts of M o r o c c o , w h i c h will therefore not concern us
f u r t h e r in this v o l u m e , occurs in semiarid regions. Of
c o n c e r n to us is a pitcher plant, Sarracenia purpurea,
which can occasionally be found in alkaline marl bogs
of the n o r t h e a s t as well as in its usual h o m e in acid
bogs.
N o w we are going to m a k e a quick tour of major
sections of N o r t h A m e r i c a and take an overview of
b r o a d areas w h e r e categories of carnivorous plants can
be f o u n d in suitable locations within the various areas.

Fig. 1-3. Sarracenia purpurea, a pitcher plant, with a tubu-

lar leaf trap of the pitfall type.

Fig. 1-4. Drosera capillaris, a sundew, with traps of the

"flypaper," or adhesive, type. The entire flattened rosette
is about 5 cm across. Note the numerous gland hairs with
sticky secretions at their tips.

6 / C a r n i v o r o u s Plants of the United States and Canada

 In the eastern t w o - t h i r d s of C a n a d a and the n o r t h - purpurea in some of these areas indicates a n o t h e r f o r m
eastern quadrant of the U n i t e d S t a t e s , carnivorous of a d a p t a t i o n that is not completely u n d e r s t o o d . A
plants are m o s t o f t e n f o u n d in the classic acid s p h a g - sundew, Drosera linearis, is adapted to marl bogs
num bog with w h i c h even the w e e k e n d naturalist is around the G r e a t L a k e s , where it is f o u n d a l m o s t e x -
likely familiar. A n o r t h e r n s p h a g n u m b o g is usually clusively. O n the other h a n d , m a n y o t h e r c a r n i v o r o u s
an ancient glacial lake that has m a t u r e d into a b o g p l a n t s will n o t colonize marl b o g s . O n e final p o i n t on
by b e c o m i n g partially filled with undecayed plant the e c o l o g y of the n o r t h e a s t e r n region is that it h a s
detritus. It is then overgrown by large m a s s e s of v a r i - b e e n repeatedly glaciated, and a f t e r e a c h ice flow r e -
ous species of Sphagnum and other m o s s e s , all tending treated, plants h a v e m o v e d n o r t h again to repopulate
to produce a very acid growing m e d i u m . As the b o g suitable sites. T h u s , plant populations in this region
further m a t u r e s , or ages, o t h e r small p l a n t s , followed h a v e b e e n stable for relatively short geologic periods.
by larger woody plants, gradually m o v e in toward the P r o g r e s s i n g a little s o u t h , we c o m e to the r e m a r k a b l e
center of the former lake until finally a n o r t h e r n forest N e w J e r s e y Pine B a r r e n s , w h i c h is a g r o s s m i s n o m e r
results. Carnivorous and o t h e r b o g plants are t h e n since to the eye of a naturalist t h e y are a n y t h i n g but
crowded out. b a r r e n . But the early colonists did not find the b r o a d ,
But while the bog is relatively y o u n g , one can o f t e n s w e e p i n g , sandy hills conducive to f a r m i n g , so they
find the pitcher plant Sarracenia purpurea g r o w i n g in declared t h e m b a r r e n , and only timber and m i n i n g
profusion along with various s u n d e w s (Drosera) and interests were able to utilize the region to a n y extent.
some bladderworts (Utricularia), the latter either in Here there are m a n y acid b o g s along and in old l a k e s ,
the sphagnum or in the open acid w a t e r o f t e n f o u n d slow s t r e a m s , and sluggish springheads. T h e pitcher
in the center of a bog. M a n y b o g s are still basically plant Sarracenia purpurea is quite abundant, and the
large lakes or ponds with more open w a t e r t h a n s p h a g - kinds of sundews (Drosera) and b l a d d e r w o r t s (Utricu-
num m a t , and in such areas the carnivorous plants laria) become much more diverse; but butterworts
grow along the lake m a r g i n s . In m o r e s a n d y , open (Pinguicula) are absent.
places along the shores of large lakes and the G r e a t In the s o u t h e r n A p p a l a c h i a n M o u n t a i n chain f r o m
Lakes, b u t t e r w o r t s (Pinguicula) m a y be f o u n d . P e n n s y l v a n i a south to its terminus in A l a b a m a , there
We mentioned briefly the marl b o g . T h i s is a special are occasional relic b o g s that have survived ancient
area in which the seepage of spring water over a flat geologic activity that created these m o u n t a i n s f r o m a
surface causes calcium c a r b o n a t e to percolate up f r o m peneplain (an a l m o s t level p l a i n ) . T h e b o g s are v e r y
limestone deposits. T h e alkaline marl results in c o n - m u c h like the acid s p h a g n u m t y p e of the f a r n o r t h
ditions just the opposite f r o m those of the s p h a g n u m in general a p p e a r a n c e , b u t t h e y are m o s t o f t e n f o u n d
bog. But marl b o g s do otherwise have some of the f e a - at a confluence of springheads or beside a s t r e a m r a t h e r
tures that allow the g r o w t h of carnivorous p l a n t s , than around the edges of maturing glacial l a k e s , w h i c h
among them diminished n i t r o g e n and o t h e r salts, c o n - are n o t present in these areas. T h e kinds of carnivorous
stantly wet conditions, and the a b s e n c e of m a n y o t h e r p l a n t s f o u n d in these m o u n t a i n b o g s are l i m i t e d : the
plants that might b e c o m e competitors. T h e presence of pitcher plant Sarracenia purpurea, a sundew (Drosera),
the normally acid-loving pitcher plant Sarracenia a f e w bladderworts (Utricularia), and t w o o t h e r pitcher

7/ An Introduction
 plants that are unusual cases and will be discussed in
C h a p t e r 3. M o r e b o g s are found as the mountains r e -
cede into the eastern foothills and piedmont sections of
the s o u t h e a s t e r n s t a t e s , and the diversity of carnivo-
rous plants increases as we approach the coast.
T h e last general area to consider in the east is the
s o u t h e a s t e r n coastal plain, which runs as a great arc
f r o m eastern Virginia south and west to eastern T e x a s ,
including all of Florida. T h i s area was suboceanic b e -
f o r e the coastal uplift, and it is p r o b a b l e that the rich
carnivorous plant life there is ultimately descended
f r o m plants of the f o r m e r peneplain which has n o w
b e e n replaced b y m o u n t a i n s and piedmont. T h e plants
apparently migrated d o w n rivers to habitats more simi-
lar to their ancestral locations. Since then, further cross
m i g r a t i o n s and evolution have undoubtedly occurred.
T h e f e w f o r m s adaptable to m o u n t a i n climates were
able to stay b e h i n d and evolve still f u r t h e r , some p r o b -
ably not adapting at all to coastal h a b i t a t s , and some
adapting to b o t h m o u n t a i n s and plain.
T h e s o u t h e a s t e r n coastal plain is our richest area for
b o t h the n u m b e r of species and the total population
of carnivorous p l a n t s ; a b o u t ninety per cent of the
species to be discussed in this b o o k can be found there.
M a n y , such as the V e n u s ' flytrap (Dionaea muscipula),
are f o u n d there exclusively. W h i l e s p h a g n u m b o g s of
the streamside or springhead type are found in this

Fig. 1-5. A typical Appalachian Mountain bog. Note the

background trees and mountaintop. The bog is grassy with
a ground layer of sphagnum moss. There is water 2 - 3 cm
deep in most places. This is a confluent spring bog, and the
drainage stream is seen in the lower right-hand corner.

Fig. 1-6. A southeastern coastal plain savannah. The trees

are lightly scattered among grasses and sedges. Some pitch-
ers of Sarracenia flava can be seen above the grass in the
foreground.

8/ Carnivorous Plants of the United States and Canada

region in a b u n d a n c e , the m o s t characteristic h a b i t a t
is a s a v a n n a h , or grass-sedge bog. T h i s is a l o w , flat or
slightly sloping, sandy area with high w a t e r table and
supporting p r e d o m i n a n t stands of g r a s s e s , sedges, and
rather widely spaced longleaf pines. A h e a l t h y s a v a n -
nah is quite m o i s t and acid.
T r a v e l i n g rapidly across the c o n t i n e n t , there is a
paucity of carnivorous plants in the mid-plains and
prairie: one species of sundew (Drosera) in wet p o c k e t s
of the southern plains, and some bladderworts (Utricu-
laria) in scattered aquatic sites. T h e deserts and eastern
R o c k y M o u n t a i n s are devoid of carnivorous plants.
Parts of the Pacific m o u n t a i n slopes are a different
matter, particularly f r o m n o r t h e r n C a l i f o r n i a into O r e -
gon. A g a i n there are s p h a g n u m b o g s alongside or
heading mountain s t r e a m s , as there are in the eastern
mountains. C u r i o u s l y , there are several of the same
kinds of sundew (Drosera) and bladderworts (Utricu-
laria) that are f o u n d in the east, and a b u t t e r w o r t
(Pinguicula) can also be found there. Q u i t e unique is
the California pitcher plant, Darlingtonia californica,
a m e m b e r of the same family ( S a r r a c e n i a c e a e ) as the
eastern pitcher plants but ranked in its o w n genus.
THE FUTURE OF C A R N I V O R O U S PLANTS
T h i s section title propounds a vital question to w h i c h
we can offer only some g u e s s e s , but t h e y are largely
well-founded guesses. Unfortunately, the outlook
appears quite grim for m a n y species. A n u m b e r of
factors contribute to this opinion.
Primarily, the m o s t dangerous hazards are the r e -
sult of m a n ' s modification of the e n v i r o n m e n t for p e r -
sonal and often shortsighted ends. T h e s e modifications
include, particularly, the control of fire and w a t e r
levels in wetlands.
We will first consider the value of fire. T h e s u g g e s -
tion that there is any " v a l u e " in fire would seem a n a t h -
9/An Introduction
e m a to t h o s e w h o in the past c e n t u r y h a v e put all
the efforts and sloganeering into the control of fire
t h r o u g h o u t our wild lands. But w h a t we will be talking
a b o u t is a specific kind of fire in certain specific areas.
Fire is n e c e s s a r y f o r the health of a b o g . B o g core
samples s h o w that in ancient times there were m a n y
fires, as is indicated by charcoal layers a n d evidence
of post-fire r e g r o w t h . M o d e r n research in w h i c h b o g s
h a v e b e e n regularly fired o v e r a period of years s h o w s
that a f a s t surface fire tends to remove detritus, c o m -
peting h e r b s , and y o u n g w o o d y plants that invade the
m a r g i n s of a b o g as it goes t h r o u g h its natural m a t u r a -
tion process (eutrophication) toward b e c o m i n g forest
land in the n o r t h and a scrub b o g in the south. Periodic
a u t u m n firing, properly c o n t r o l l e d , can greatly prolong
the life of a b o g . H o w e v e r , m a n the f a r m e r , l u m b e r -
m a n , and developer has either caused superheated
h o l o c a u s t s that destroy e v e r y t h i n g in huge tracts of
natural lands or has tried to control all fire. M a n y fine
areas f o r m e r l y i n h a b i t e d by carnivorous plants have
b e e n a b s o r b e d into forest or scrub during the lifetimes
of living b o t a n i s t s w h o have witnessed the process.
S e c o n d , in order to be able to approach t i m b e r l a n d s
in s w a m p s , as well as to e x t e n d agricultural a r e a s ,
m a n y wetlands h a v e b e e n and still are being drained.
O t h e r s have b e e n converted to p o n d s or lakes. In the
relatively flat terrain of the s o u t h e a s t e r n coastal plains,
w h e r e an area of several hundred acres m a y not h a v e
a variance in net elevation of m o r e than a few f e e t ,
drainage is easily accomplished and can be clearly seen
in the n e w l y created p a t c h e s of seasonal desert amid
an e x t e n s i v e n e t w o r k of roadside ditches t h r o u g h o u t
this section of the c o u n t r y .
T o these t w o main factors can b e added such s e c o n -
dary insults as pollution with fertilizer and toxic m a -
terials, willful v a n d a l i s m , and the collection of p l a n t s
b y casual e n t h u s i a s t s w h o are passing t h r o u g h . T h e r e
h a v e even b e e n recent d o c u m e n t e d instances of the
collection by commercial nurseries of entire stands of
extremely rare f o r m s . T h e s e p r o b l e m s do not eclipse
the more basic situation of a radical c h a n g e in the
h a b i t a t , but they are not at all m i n o r , the excesses of
h u m a n n a t u r e being w h a t they are.
So m u c h f o r the g r i m side of the picture. On the
o t h e r h a n d , there are ongoing a t t e m p t s to preserve
representative areas f r o m the fate of n e i g h b o r i n g l o -
cations. T h e s e efforts are having v a r y i n g success. M o r e
people are b e c o m i n g seriously c o n c e r n e d a b o u t the
misappropriation of our r e s o u r c e s , even those resources
in which no immediate e c o n o m i c value is apparent.
N a t u r e c o n s e r v a n c i e s , provincial, state, and n a t i o n a l
parks and a f e w local o n e s , local private groups p r e -
serving a small b o g , and b o t a n i c a l gardens featuring
native plants are all m a k i n g some h e a d w a y in setting
aside, p r o t e c t i n g , and m a n a g i n g wisely areas that in-
clude carnivorous plants. S o m e states have passed
sweeping plant protection l a w s , a l t h o u g h e n f o r c e m e n t
of these laws is difficult and at b e s t erratic. S o m e c o m -
mercial l u m b e r c o m p a n i e s in the s o u t h e a s t e r n U n i t e d
S t a t e s h a v e , on the r e c o m m e n d a t i o n of experienced
b o t a n i s t s , t a k e n upon themselves the task of sparing
and even annually b u r n i n g certain b o t a n i c a l l y valuable
tracts of land w h i c h could h a v e b e e n devoted to tree
f a r m i n g . H o w e v e r , these same c o m p a n i e s grant " c a r -
n i v o r o u s plant collection and sales r i g h t s " to c o m m e r -
cial nurseries.
T h e r e is a lot to be d o n e , and the situation is rather
urgent. No individual is going to be able to m a k e d r a -
m a t i c c h a n g e s o f a n y sort, and m u c h o f the d a m a g e
is irreversible, short of reclaiming sites t h r o u g h radical
t r e a t m e n t and then m a k i n g m a s s i v e t r a n s p l a n t i n g s .
But individuals can participate in and support c o n s e r -
vation groups w h i c h are t r y i n g to set aside a f e w
extant representative a r e a s , w h e t h e r t h e y are areas of
1 0 / Carnivorous Plants of the United States and Canada
p a r k proportions or small b o g s located on local f a r m -
land w h i c h m i g h t be purchased and properly m a i n -
tained. On an individual basis, one can pursue one's
citizen's rights by electing sympathetic legislative r e p -
resentatives or influencing the votes of those already
elected, keeping in mind the realistic fact that a certain
proportion of desirable natural lands must and will
yield to basic e c o n o m i c and h u m a n necessity. Also on
an individual b a s i s , one can discourage vandalism and
suppress o n e ' s o w n inclination to dig and try growing
unusual native plants alongside the tomatoes and p e -
tunias. T h e y will always die with such casual treatment.
T h e serious student can be of further help by assist-
ing in preserving in artificial or barren natural b o g s ,
in t u b s , or in g r e e n h o u s e s , m a n y species of carnivorous
plants that are collected f r o m condemned sites or pur-
chased t h r o u g h reputable dealers. D e a l e r s are supposed
to p r o p a g a t e their s t o c k rather than collect f r o m the
field to fill orders. A few reliable commercial sources
f o r carnivorous plants are given at the conclusion of
this b o o k , along with hints on growing the plants
successfully.
T h e last t w o sections of this chapter are intended
f o r the reader w h o is not widely experienced b o t a n i -
cally. T h e y are very brief reviews of flower structure
and f u n c t i o n and of the s y s t e m of Latin biological
n a m e s . T h o s e desiring additional information should
consult any e l e m e n t a r y b o t a n y text or some of the ref-
erences m e n t i o n e d in the final chapter.
FLOWER STRUCTURE
T h e flowers of m o s t carnivorous plants take second
place in attention to the trap leaves. But in m a n y b l a d -
derworts (Utricularia), the flowers may be the most
n o t i c e a b l e part of the plant and the part m o s t easily
used f o r identification. Flowers are, of course, impor-
tant reproductive o r g a n s , and some familiarity w i t h
their general plan is n e c e s s a r y in any b o t a n i c a l study.
Reproduction is obviously f u n d a m e n t a l to all living
organisms. T h e usual results of wear and tear, age, and
diseases would soon b e c o m e self-evident if plants did
not continually replace their losses. W e m a y b r o a d l y
divide reproduction into (1) sexual, the e x c h a n g e of
living material b e t w e e n two separate o r g a n i s m s of the
same kind so as to diversify and e n h a n c e the genetic
base of the species, and (2) asexual, the division of
parent plants or parts of p l a n t s , which allows rapid
local reproduction under sometimes adverse c i r c u m -
stances. Asexual m e t h o d s of reproducing include the
familiar bulbs, rhizomes, s t o l o n s , and various types of
budding. M a n y plants utilize b o t h general categories
of reproduction. T h e flower is the organ of sexual r e -
production in green seed-bearing plants.
11/An Introduction
M o s t flowers consist of four basic whorls or layers
of structures w h i c h , f r o m b o t t o m to top (or outside
i n ) , are (1) sepals, collectively called the calyx, w h i c h
are usually green and leaf like but can be c o l o r f u l ;
(2) petals, collectively called the corolla, which are
usually quite colorful and o f t e n give the flower the
general a p p e a r a n c e by w h i c h it is recognized; (3)
stamens, each consisting of a simple filament s u p p o r t -
ing a k n o b l i k e b o d y called the anther, which actually
produces the pollen, a fine dusty m a t e r i a l ; and (4) the
pistil, at the top of w h i c h are one or m o r e stigmas
having s t i c k y , s u g a r y secretions that hold the pollen
grains and support their germination. Supporting the
stigma is the filamentous style, below which is located
the enlarged, o f t e n b u l b o u s ovary, which houses the
egg cells and will ultimately b e c o m e the seed capsule.
S o m e flowers have eliminated the calyx, the corolla,
or b o t h , and some species have either stamens or pis-
tils but not b o t h . N o n e of these conditions occur in
the N o r t h A m e r i c a n carnivorous plants, h o w e v e r .
T h e entire flower is supported on a base called a
peduncle, and the whole is atop a scape, or flower
"stalk."
S o m e t i m e s , if there is more than one flower to a
scape, another stemlike structure called a pedicel
c o n n e c t s the flower b a s e to the scape. T h e latter c o n -
dition is especially p r o m i n e n t a m o n g bladderworts
(Utricularia).
T h e sexual aspect o f plant reproduction, the e x -
c h a n g e of living material b e t w e e n two plants of the
s a m e kind or b e t w e e n compatible plants of two dif-
f e r e n t but closely related k i n d s , ideally t a k e s place
w h e n pollen f r o m one plant is deposited on the stig-
ma of a n o t h e r . T h i s process is called cross-pollination.
Pollination m a y t a k e place with the aid of g r a v i t y ,
w i n d , w a t e r , or visiting insects or other animals t r a n s -
porting pollen f r o m one flower to a n o t h e r . A l t h o u g h
cross-pollination is conducive to the greatest benefits o n s , p e a s , m o s t o r c h i d s , a n d , a m o n g carnivorous plants,
of sexual r e p r o d u c t i o n , m a n y plants are capable of self- the bladderworts (Utricularia) and the butterworts
pollination if crossing c a n n o t occur. W h i l e sexual in a (Pinguicula).
partial s e n s e , s e l f - p o l l i n a t i o n , with respect to genetic
e x c h a n g e , is definitely inferior to crossing. Selfing,
h o w e v e r , is still superior to asexual types of reproduc-
tion, since certain biological p r o c e s s e s occurring in p o l -
lination and seed f o r m a t i o n still allow some m e a s u r e
o f genetic variation and r e c o m b i n a t i o n w h i c h c a n n o t
be achieved in vegetative reproduction.
If the deposited pollen grain h a s f o u n d a suitable
medium in the s t i c k y , s u g a r y secretions of the s t i g m a ,
it will g e r m i n a t e s o m e w h a t like a tiny seed. A pollen
t u b e carrying o n e or m o r e pollen nuclei actually g r o w s
d o w n t h r o u g h the supporting style into the o v a r y ,
w h e r e a pollen nucleus unites w i t h an egg-cell n u -
cleus. T h i s process is called fertilization. A c o m p l e x
series of microscopic cellular divisions t a k e s place in
order to b a l a n c e the nuclear genetic m a t e r i a l , and an
Actinomorphic Flower Form
e m b r y o i s f o r m e d . A r o u n d the e m b r y o , nutritional m a -
terial o f t e n develops along w i t h a seed coat, and the Radially Symmetric
ovary b e c o m e s a m a t u r i n g seed capsule w h i c h opens
w h e n ripe. T h e seeds are t h e n dispersed through vari-
ous m e a n s .
O n e additional pair of definitions needs to be c o n -
sidered. In b a s i c f o r m , flowers m a y be of two k i n d s :
(1) actinomorphic, or radially s y m m e t r i c a l . If the flow-
ers are cut in h a l f along any plane that passes through
the c e n t e r of the flower, t w o equal m i r r o r - i m a g e halves
will a l w a y s result. In o t h e r w o r d s , these are the " p e r -
fect c i r c l e " flowers, such as day lilies, m a g n o l i a s , p i n k s ,
a n d , a m o n g carnivorous p l a n t s , the V e n u s ' flytrap
(Dionaea), the pitcher plants (Sarracenia), and the s u n -
dews (Drosera). (2) zygomorphic, or bilaterally sym-
metrical. T h e s e flowers can be cut in only o n e plane
Zygomorphic Flower Form
through the center in order to o b t a i n t w o equal m i r r o r -
image h a l v e s . E x a m p l e s of such flowers are s n a p d r a g - Bilaterally Symmetric

12 / Carnivorous Plants of the United States and Canada

H O W PLANTS ARE NAMED
T h e proper n a m i n g of plants has a l o n g , c o m p l e x ,
and interesting h i s t o r y , w h i c h we will not go into h e r e .
We will give only the m o s t i m p o r t a n t principles of i n -
troductory n o m e n c l a t u r e so that the reader can find his
way in this or any basic biology b o o k . We hope to o v e r -
come the initial reaction of recoil at the sight or sound
of Latinized biological n a m e s .
T h e Latin n a m e s that b o t a n i s t s use represent a r a t h e r
successful attempt to bring a b o u t u n i f o r m i t y in n a m i n g
plants and to be certain that the s a m e n a m e is n o t used
f o r two or more plants. Lately, they tend also to reflect
theories as to various relationships b e t w e e n plants with
respect to evolution and f u n c t i o n . Latin n a m e s take
some getting used t o , but once one begins to u n d e r s t a n d
them and to associate t h e m with real entities, t h e y b e -
come easier to deal with and their a d v a n t a g e s o v e r
c o m m o n n a m e s are appreciated. T h e y are certainly here
to s t a y ; they are e n j o y i n g m o r e c o m m o n use a m o n g
n o n p r o f e s s i o n a l s , and they are the only n a m e s that are
accepted officially.
C o m m o n or local " E n g l i s h " n a m e s suffer f r o m a lack
of specificity, a c c u r a c y , and u n i f o r m i t y b e c a u s e of local
history and custom. T h u s , the c o m m o n n a m e " p i t c h e r
p l a n t " is incomplete in that it could r e f e r to any of
eight different kinds of plants or to all eight as a unit,
even though the speaker m a y have a specific plant in
mind. In another locality, the same pitcher plant could
be called a yellow t r u m p e t , a flycatcher, or a lily. In
still other areas, the term " f l y c a t c h e r " could refer to
another of the several kinds of pitcher p l a n t s , or even
to another class of carnivorous plants altogether. In
other w o r d s , c o m m o n n a m e s f r e q u e n t l y do n o t have
universal application.
The keys to eliminating this confusion revolve
around two f a c t o r s : the standardized basic Latinized
binomial s y s t e m , which has been in use f o r three h u n -
13/An Introduction
dred y e a r s , a n d the results of several recent i n t e r n a -
tional c o n g r e s s e s o f b o t a n i s t s , w h o decided h o w the
s y s t e m should be used and w h e n and h o w it should be
modified. T h e resulting rules are f a r f r o m p e r f e c t , b u t
one can e s t a b l i s h official n a m e s — o n e f o r e a c h kind of
p l a n t — a n d these n a m e s will be used and recognized
b y legitimate b o t a n i s t s the world over. T h e s a m e n a m e
c a n n o t be u s e d officially f o r any kind of plant o t h e r
t h a n that f o r w h i c h it w a s intended. T h e r e results a
c o m m o n r e f e r e n c e b a s e and a classification that sweeps
aside m u c h of the c o n f u s i o n that would be involved in
using c o m m o n n a m e s alone.
T h e b a s i c Latinized n a m e o f any plant consists o f
t w o parts o r w o r d s : the g e n u s n a m e first, followed b y
the species n a m e . T h e g e n u s n a m e i s s o m e w h a t anal-
ogous to the h u m a n s u r n a m e , and the species n a m e
c a n b e c o m p a r e d t o the " g i v e n " n a m e identifying a n
individual in the h u m a n f a m i l y . ( A c t u a l l y , the species
n a m e identifies a w h o l e population of like plants.) All
the m e m b e r s , or species, of a genus are similar to e a c h
o t h e r in s o m e w a y s , as is true in h u m a n f a m i l y r e s e m -
b l a n c e s , y e t t h e y are sufficiently different to w a r r a n t
their species n a m e s . As a g r o u p , one g e n u s is quite dif-
ferent f r o m a n o t h e r , b u t even a m o n g a n a s s o r t m e n t o f
g e n e r a (plural o f g e n u s ) there m a y b e some b a s i c
similarities and some groups of genera thus can be
b l o c k e d into b o t a n i c a l families. T h e r e are even higher
divisions, right up to that b e t w e e n the animal and plant
k i n g d o m s , b u t t h e y need not c o n c e r n us at this time.
As an e x a m p l e of classification by f a m i l y , g e n u s , and
species, let us m e n t i o n the pitcher plants of eastern
North America. T h e y are m e m b e r s o f t h e family
S a r r a c e n i a c e a e , w h i c h includes three genera of plants
f r o m N o r t h and S o u t h A m e r i c a . T h e g e n u s n a m e for
the eastern N o r t h A m e r i c a n pitcher plant is Sarracenia,
w h i c h in this case as in m a n y is very similar to the
f a m i l y n a m e since it is felt to be the prime e x a m p l e (or
type genus) of the f a m i l y . W i t h i n the g e n u s Sarracenia
there are eight widely recognized species, including
Sarracenia flava, Sarracenia purpurea, Sarracenia alata,
etc. T h u s the b o t a n i s t can s p e a k of a f a m i l y by its
proper n a m e w h e n discussing a very b r o a d group of
s o m e w h a t similar p l a n t s , a g e n u s w h e n he wishes to
m e n t i o n m o r e closely related plants as a subgroup
within that f a m i l y , and use a binomial n a m e ( o f t e n
called j u s t " s p e c i e s " in j a r g o n ) w h e n discussing a single
kind of plant.
We m u s t n o w m e n t i o n a f e w simple rules as e s -
tablished by the b o t a n i c a l congresses devoted to this
p r o b l e m . T h e n a m e of a g e n u s or species is ideally a
Latinized description of a distinctive feature of the
p l a n t , but it c a n be derived f r o m a n o t h e r source such
as a p e r s o n ' s n a m e or a g e o g r a p h i c a l location. T h e
genus n a m e a l w a y s b e g i n s with a capital letter and the
species n a m e w i t h a small letter, except w h e n the
species is n a m e d a f t e r a p e r s o n , in w h i c h case a capital
letter may be u s e d , b u t the trend is a w a y f r o m all
capitalization in species n a m e s . A binomial n a m e is
always printed in italics or underlined w h e n h a n d w r i t -
ten or typed.
In very f o r m a l b o t a n i c a l w r i t i n g , the binomial n a m e
is followed in ordinary r o m a n t y p e by the n a m e of the
person w h o first n a m e d the p l a n t , or by several n a m e s
in some cases when the n a m e h a s b e e n properly
c h a n g e d o v e r a period of time. M o s t o f t e n , a s t a n -
dardized a b b r e v i a t i o n of the m a n ' s n a m e is used if he
is well k n o w n a m o n g b o t a n i s t s .
A g e n u s n a m e can be used o n l y o n c e in b o t a n y , and
no t w o kinds of plants w i t h i n a g e n u s can bear the
s a m e species n a m e . A s a n e x a m p l e o f the latter, our
Sarracenia g e n u s of p i t c h e r p l a n t s c a n h a v e o n l y one
species named Sarracenia purpurea, but the species
n a m e purpurea can be used in o t h e r plant g e n e r a , as in
the case of Utricularia purpurea. Following the rules,
14 / Carnivorous Plants of the United States and Canada
there is no Sarracenia o t h e r than one of the eastern
N o r t h A m e r i c a n pitcher plants.
A n o t h e r rule is that certain abbreviations are allow-
able. If one is discussing Sarracenia in a writing, one
c a n a b b r e v i a t e the g e n u s to the first letter after once
using the full genus n a m e . S o , if I mention Sarracenia
flava in a chapter or paragraph and have not during
the course of the discussion mentioned another genus
b e g i n n i n g w i t h the letter S, I m a y abbreviate the
n a m e to S. flava the n e x t time I use it.
N o t all is settled by any m e a n s in plant naming and
classification (the science and art of t a x o n o m y ) , and
we will have to c o n f r o n t a few controversies in this
b o o k . T h e r e is o f t e n debate a b o u t which n a m e has
legal priority in a case where two botanists have acci-
dentally or willfully given the same kind of plant
t w o different Latin n a m e s . T h e r e is m u c h serious and
legitimate discussion about w h a t actually constitutes
a s p e c i e s — w h e r e one draws the line b e t w e e n " k i n d s "
of plants. T h e g e n u s Sarracenia has eight c o m m o n l y
accepted species, b u t m a n y serious and learned b o t a -
nists with good a r g u m e n t s would declare that there
are as m a n y as ten species. H o w e v e r , in order to get
additional species established they would have to go
t h r o u g h the procedures that have been prescribed for
such c h a n g e s , and all b o t a n i s t s would have access to
the i n f o r m a t i o n .
T h e c o m p l e t e classification of a w h o l e group of
p l a n t s , or a single species, can be changed if certain
f o r m a l steps with respect to studies and the publica-
tion of those studies are carried out, and t h e n — m o s t
importantly—if enough botanists agree with those
studies and use the n e w classification. W h e t h e r they
do or n o t , all b o t a n i s t s would k n o w precisely w h a t w a s
being discussed if t h e y came across a paper on the s u b -
j e c t . A d m i t t e d l y , as in the case of rules and laws
e v e r y w h e r e , a considerable a m o u n t of nonformalized
agreement is necessary to m a k e the s y s t e m w o r k well. stones of evolutionary c h a n g e . At the very least, their
A n o t h e r area of c o n t r o v e r s y in t a x o n o m y is the use value lies in their u s e f u l n e s s in c o m m u n i c a t i o n , if
of subdivisions finer than species, these b e i n g , in order n o t h i n g else. T h e horticulturist can find a varietal n a m e
of specificity, s u b s p e c i e s , f o r m s , varieties, and races. very helpful f o r accurate and b r i e f r e f e r e n c e w h e n
A few subspecies, representing differences in large communicating with other horticulturists, and the
populations of a species that occur over wide areas or ecologist or evolutionist studying the derivation and
with discontinuous r a n g e s , are f o r m a l l y but reluctantly relationship of plants can find the subdivisions equally
recognized by m a n y students. Forms and varieties are useful for quick and pointed reference to natural v a r i a -
more or less tentative and most o f t e n represent p o p u l a - tions that he m a y recognize for one reason or a n o t h e r .
tions of more importance to the ecologist and the e v o - Proper a b b r e v i a t i o n s f o r handling these subdivi-
lutionist than to the taxonomist. O f t e n they are merely sions are ssp. ( s u b s p e c i e s ) , f. ( f o r m a ) , and var. or v.
horticultural terms, m a n y of w h i c h are eventually ( v a r i e t y ) . T h e subdivisional n a m e is italicized, but
dropped altogether, but the plants they represent m a y the r e f e r e n c e a b b r e v i a t i o n is n o t . Examples are Sarra-
after further study be elevated in stature to subspecies cenia purpurea ssp. venosa and Drosera filiformis f.
o r s p e c i e s — a l t h o u g h this h a p p e n s rarely. M a n y t a x o n - tracyi. O f t e n the divisional abbreviation is o m i t t e d , as
omists claim that these finer subdivisions are artificial, in Sarracenia purpurea venosa. We will, as we go
a product of m a n ' s propensity f o r organization rather a l o n g , m e n t i o n i n s t a n c e s of such subdivisions a m o n g
than representing actual biological groupings. O t h e r s carnivorous p l a n t s and some of the pertinent a r g u -
claim that the variants they represent are the stepping- m e n t s pro or con.

15 / An Introduction
II. The Venus' Flytrap (Dionaea muscipula)
BOTANICAL NAME: Dionaea muscipula Ellis ex L.
A m o n o t y p i c g e n u s ; that is, there is only o n e species
in the g e n u s . Family D r o s e r a c e a e .
C O M M O N N A M E S : V e n u s ' f l y t r a p , f l y c a t c h e r , tip-
itiwitchet, catch-fly sensitive. ( T h e latter t w o n a m e s
are ancient.)
R A N G E : Q u i t e localized i n scattered s a v a n n a h s o f
s o u t h e a s t e r n N o r t h C a r o l i n a and n e i g h b o r i n g eastern
S o u t h C a r o l i n a in an a p p r o x i m a t e landward radius of
6 0 - 7 5 miles a r o u n d W i l m i n g t o n , N . C .

Fig. 2-1. A plant of Dionaea muscipula as typically seen in

FLOWERING SEASON: Late May through early the field.
June.
TRAP SEASON: S o m e plants remain evergreen i n enlarges a n n u a l l y . Fibrous roots descend 1 0 - 1 5 c m .
protected s i t u a t i o n s , while m a n y die b a c k f o r the T h e green leaves g r o w up to 12 cm long. T h e y are of
w i n t e r . N e w traps begin g r o w i n g i n M a r c h and c o n - two p a r t s : a n a r r o w to relatively b r o a d leaf like petiole
tinue into O c t o b e r . (leaf stem) n e a r e r the rosette c e n t e r , and a leaf blade
modified into a unique t r a p , m e a s u r i n g up to 3 cm long.
D E S C R I P T I O N . — T h e plant is a rosette of leaves that - T h e flowers are on a 1 - 3 0 cm scape. T h e y are acti-
radiate out f r o m a central p o i n t , the leaves being totally n o m o r p h i c , h a v e white p e t a l s , and are about 1 . 0 - 1 . 5
o r partially reclining. T h e rosette m e a s u r e s 1 0 - 1 4 c m c m a c r o s s . A f t e r fertilization, t i n y , b l a c k , pear-shaped
across w h e n m a t u r e . T h e leaves arise f r o m a s o m e w h a t seeds set (mature) in 6 - 8 w e e k s . T h e s e will germinate
e l o n g a t e , f l e s h y , w h i t e r h i z o m e (underground stem) immediately when sown on a suitable substrate;
o f t e n miscalled a " b u l b . " T h e rhizome e l o n g a t e s and storage at w a r m t e m p e r a t u r e s results in a lower rate

16 / Carnivorous Plants of the United States and Canada

of germination. T h e plants m a t u r e f r o m seedlings to
flowering age in 3 - 4 y e a r s , and estimates of the age of
the oldest k n o w n living plants are nearly 25 y e a r s .
The traps consist of two clamshell-like halves.
Around the free m a r g i n s (unattached edges) are n u -
merous stout guard hairs and minute n e c t a r glands.
T h e trap is normally in a 4 5 ° - 6 0 ° open position w h e n
undisturbed. T h e interior of each half is lined by n e a r -
ly microscopic digestive glands, which give the surface
a finely granular or c o b b l e s t o n e d appearance. Each
inner half has also three smaller, finer trigger hairs in
a triangular pattern (rarely a n o m a l o u s plants have up
to six hairs per half) w h i c h , w h e n properly stimulated,
initiate trap closure. T h e trap lining is colored v a r i a b l y
green to pale yellow to bright red with f r e q u e n t i n t e r -
mediate shades and patterns noted.
T h e usually bright coloration and the secretions of
sweet nectar by the marginal glands m a y attract prey
to the interior of the t r a p , w h e r e the insect b r u s h e s
against one or more of the trigger hairs. Initially, trap
closure is quite rapid until the guard hairs m e s h , e f -
fectively incarcerating the small prey in a barred
sarcophagus. T h e slower, s e c o n d a r y p h a s e o f closure
results in the m a r g i n s sealing tightly together so that
the whole trap b e c o m e s a flattened, s t o m a c h l i k e pouch.
At this stage, the m a r g i n s of the trap halves evert
slightly. If live p r e y — n o t a raindrop or a piece of
w i n d b l o w n d e b r i s — h a s b e e n c a u g h t , digestive f l u i d s
are then secreted into the interior of the closed trap.
A p p a r e n t l y , the struggles of the prey and certain

Fig. 2-2. A plant of more upright habit. Note the numerous

red-lined traps with prominent marginal hairs.

Fig. 2-3. Close view of two traps, one with previously di-
gested insect remains, the other after the rapid phase of
closure. The intef/neshing guard hairs hold the prey in until
the slower closing phase is completed.

17 / The Venus' Flytrap

 chemical c o m p o u n d s that e m a n a t e f r o m the prey (such
as amino acids) stimulate more copious secretions of
digestive fluid with greater c o n c e n t r a t i o n s of enzymes.
D i g e s t i o n t h e n o c c u r s , and the nutrients are absorbed
at the b a s e s of the glands o v e r a period of 3 - 5 d a y s ,
depending on the t e m p e r a t u r e , the size of the prey,
its nutritive value, etc. A f t e r w a r d s , the trap reopens.
T h e d r y , chitinous remains of the insect stay in the
trap or drop out.
Each trap m a y be stimulated mechanically by touch
to close a b o u t ten times b e f o r e it will no longer re-
s p o n d . A f t e r such closures, the trap reopens the n e x t
d a y , since there is no animate matter to digest. If the
prey is n o t too large, each trap may catch and digest up
to three times, a f t e r which it ceases to function. V e r y
large catches result in the death of the trap leaf, but
n e w ones g r o w more or less continuously all season.
In order f o r the trap to close, any one of the trigger
hairs on the inner surface m u s t usually be touched
twice, or any t w o hairs m u s t be touched once each in
succession. W t t e m p e r a t u r e s a b o v e 4 0 ° C , h o w e v e r ,
one stimulus suffices in half the cases. T h e longer the
period b e t w e e n the t w o stimuli, the slower the closure.
T h e very quick reaction no longer occurs if the period
exceeds the range of 2 0 - 4 0 seconds. If the stimuli are
f a r t h e r apart, closure will eventually occur, but it is
e x t r e m e l y slow and multiple stimuli are required dur-
ing closure to complete the process.| It is felt that the
struggle of the live prey inside the trap continuously
stimulates the closure m e c h a n i s m and the secretion of

Fig. 2-4. Interior of trap. Note the three triangularly spaced

trigger hairs on each inner surface of the trap.

Fig. 2-5. Appearance of trap after catching a live insect,

the slow phase of closure having been completed. The edges
are tightly sealed.

1 8 / Carnivorous Plants of the United States and Canada

the digestive enzymes until the very tight, pouchlike
seal is formed.
T h e m e c h a n i s m of closure is not entirely understood.
A small but consistently m e a s u r a b l e action potential,
or electric current, crosses the leaf a f t e r the trigger
hair is stimulated. S o m e t i m e s , the stimulation of parts
of the leaf o t h e r than the trigger hairs initiates the
action potential and results in closure. T h e fact that
the total n u m b e r of closures for any one trap is limited
indicates that some cell g r o w t h f u n c t i o n , metabolic
process, or b o t h , is ultimately e x h a u s t e d . T h e fact that
the n u m b e r of repeat closures is more limited if prey is
caught- each time certainly indicates that the process is
accumulative rather than e x h a u s t i v e ; that is, that p o s -
sibly the storage or metabolic m a n u f a c t u r e of some
p r o d u c t — p e r h a p s starch or p r o t e i n — m a y cause the
process of closure to b e c o m e inhibited s o o n e r than it
would with inanimate stimulation.

In spite of initial appearances, the trap does not close

like a bear trap. T h e two halves do n o t rotate on the
midrib like a hinge. W h e n open, the outer surface of
each trap half is c o n c a v e , or dished in, while the i n n e r
surfaces are bulging inwards. D u r i n g closure, these
surface c o n f o r m a t i o n s are reversed, so that the free
edges are quite suddenly b r o u g h t closer t o g e t h e r —
close enough for the strong guard hairs to i n t e r m e s h .

Fig. 2-6. A cluster of plants in flower. The scapes are dis-

proportionately long for the size of the plant, perhaps so
that the flowers are raised above the grasstops for effective
pollination, and so that the potential pollinators are not
m
more tempted by the traps farther down.

Fig. 2-7. Flowers of Dionaea muscipula.

Fig. 2-8. Typical view of Dionaea flowering in the field.

The small white flowers are just visible above the grasstops,
while the vegetative parts are partially obscured.

19 / The Venus' Flytrap

 T h i s process can be noted by close, careful observation
as well as by high-speed comparison photographs or
m o t i o n pictures. H o w e v e r , the c h a n g e in surface con-
f o r m a t i o n explains only the rapid phase of closure.
D u r i n g the slower, sealing p h a s e , no further signifi-
c a n t c h a n g e in surface c o n f o r m a t i o n is observable
e x c e p t for the eversion of the very margins, and there
is even some slight loss of the new convexity of the
outside surfaces. At this stage, some hingelike rotation
on the midrib likely occurs.
As we m e n t i o n e d , the bladelike leaf petiole has a
variable m o r p h o l o g y . In shadier locations, or in the
early spring prior to flowering, the petiole is quite thin
and wide and later ones b e c o m e thicker and narrower.
But even late in the season and in full sun, some peti-
oles of adjacent plants remain wide, while others b e -
c o m e almost stemlike or triangular in cross section.
B o t h variations can s o m e t i m e s be found in plants in a
single location. S o m e o b s e r v e r s feel that these varia-
tions are i n h e r e n t in different genetic strains of the
species, and they h a v e recognized at least four such
v a r i a t i o n s , including the two extremes and two inter-
mediates. A n o t h e r interesting observation is that the
n a r r o w e r the petiole, the m o r e erect the whole leaf
tends t o b e .
T h e color of the trap lining m a y follow a similar
p a t t e r n . G e n e r a l l y , g r o w t h in bright sunlight brings
out the brightest red color. H o w e v e r , some plants
g r o w i n g right beside the red o n e s , in the same light
and soil, remain green or yellowish or are even varie-
gated red and green.
A n o m a l i e s of m o r p h o l o g y aside f r o m that of the
petiole are also occasionally n o t e d . W e have mentioned
that the trap can h a v e up to six trigger hairs per half.

Fig. 2-9. Seed, which ripen exposed.

Fig. 2-10. Seedlings.

20 / Carnivorous Plants of the United States and Canada

A n o t h e r individual, n o n g e n e t i c variation is a double o b s e r v e d c a t c h i n g small aquatic a n i m a l s . S i n c e the u n -
trap on one petiole. An occasional flower a n o m a l y is derground stem of the V e n u s ' flytrap is well p r o t e c t e d ,
vegetative apomixis, in which the flower parts— Dionaea is among the first plants to sprout back
sepals, petals, s t a m e n s , and p i s t i l — a r e replaced b y strongly in a b u r n e d area. If the area is n o t b u r n e d
miniature plantlets w h i c h can be rooted and g r o w n to f r o m time to time (or if the water table d r o p s ) , o t h e r
normal plants. T h i s occurs m a i n l y w h e n spring w e a t h e r h e r b s , s h r u b s , and trees e n c r o a c h and quite q u i c k l y
has been uneven during the early period of flower b u d crowd out the smaller Dionaea, since the h a b i t a t is
initiation, with cold nights alternating with w a r m , s u n - t h e n a completely different o n e . T h u s a rapid s u r f a c e
ny days. T h e author produced this p h e n o m e n o n some fire in the a u t u m n is actually quite beneficial.
years ago in O h i o by growing the plants on a w i n d o w - In late s u m m e r , b e c a u s e the n e i g h b o r i n g grasses are
sill during March. The warm sunlight streaming quite tall by that t i m e , considerable search is required
through the window initiated early g r o w t h and flower by the uninitiated b e f o r e he finds the p l a n t s , often
budding, but the nights were so cold that the glass a f t e r w a l k i n g o v e r t h e m f o r some t i m e ! T h e best time
n e x t to the sill was o f t e n frosted on the inside in the to observe Dionaea is in early spring w h e n grasses
mornings. It is likely that other such e n v i r o n m e n t a l and sedges are s h o r t e r , and especially w h e n the flow-
s h o c k s , perhaps involving chemical s u b s t a n c e s such e r s , lifted by the tall scapes a b o v e the grass tops, can
as one or more of the plant h o r m o n e s , could precipitate be seen easily even f r o m an a u t o m o b i l e .
the process. Finally, even the w i d e n e d , bladelike peti- Dionaea is r a t h e r h a r d y climatically. Outdoor ex-
oles are capable of producing vegetative b u d s . T h e perimental t r a n s p l a n t s h a v e thrived as far north as
process has been utilized in culture in order to p r o p a - N e w J e r s e y and s o m e b o g s i n P e n n s y l v a n i a . Incident-
gate the plants rapidly, and the p h e n o m e n o n has o c - ally, such t r a n s p l a n t s , w h i c h w e r e o f t e n conducted
casionally been o b s e r v e d in nature. without f a n f a r e to discourage v a n d a l s , have nearly led
to some e m b a r r a s s i n g results w h e n skilled n a t u r a l i s t s ,
GENERAL.—Largely because of habitat changes, u n a w a r e o f the e x p e r i m e n t s , have c o m e u p o n the
Dionaea muscipula is m a r k e d l y decreasing in n u m b e r s plantings during w a l k s and n e a r l y rushed to publi-
throughout a shrinking range w h i c h is n o n e too large c a t i o n with the news of Dionaea's supposed range
to begin with. Indeed, earlier reports d o c u m e n t a far extension.
more extensive range in the C a r o l i n a s than we are W h i l e there is certainly m u c h m o r e to learn about
able to report n o w . Dionaea does tend to remain on all the plants in this book, Dionaea muscipula un-
a deteriorating site longer than m a n y associate c a r - doubtedly r e m a i n s the pet a m o n g m o s t students of
nivorous and noncarnivorous p l a n t s , particularly p i t c h - carnivorous p l a n t s , b o t a n i s t s in g e n e r a l , and n a t u -
er plants (Sarracenia). T h e plants g r o w in m o i s t , s a n d y , ralists of all w a l k s .
acid s a v a n n a h s a m o n g wire g r a s s e s , sedges, and m a n y
native orchids (Pogonia, Calopogon, Platanthera, Spi-
ranthes, etc.) b e t w e e n rather widely spaced longleaf
pine trees. Dionaea will tolerate short periods of
drought and flooding, s u b m e r g e d plants having b e e n

21 / The Venus' Flytrap

III. The Eastern North American Pitcher Plants
(Sarracenia L.)
The Genus
BOTANICAL NAME: Sarracenia L . N a m e d after D r .
M . S . Sarrazin o f Q u e b e c , a n early discoverer. Family
Sarraceniaceae.
COMMON NAME: Pitcher plant. (More specific
c o m m o n n a m e s will be m e n t i o n e d with each species.)
R A N G E : G e n e r a l l y , various species can b e f o u n d i n
appropriate b o g s , s a v a n n a h s , and other wet places
t h r o u g h o u t eastern N o r t h A m e r i c a .
F L O W E R I N G A N D T R A P S E A S O N S : Flowering peri-
ods vary f r o m M a r c h to J u n e , depending on species
and locale. M o s t pitchers die b a c k o v e r the w i n t e r , and
n e w ones sprout with or soon a f t e r flowering.
D E S C R I P T I O N . — P i t c h e r p l a n t s are all basically p e -
rennial rosettes of leaves modified into traps that arise
f r o m long rhizomes (underground stems) which h a v e
fibrous roots. T h e pitchers o f various species range
i n height f r o m 1 0 t o 1 2 0 c m . T h e rhizomes o f t e n b r a n c h
and have several g r o w t h c r o w n s , so that an apparent
clump of p l a n t s m a y really be a c l o n e ; that i s , the
plants m a y all be c o n n e c t e d .
T h e oldest part o f the r h i z o m e i s o f t e n dead. T w o
species regularly give rise also to flattened, m o d e r a t e l y
w i d e , e l o n g a t e , leaflike s t r u c t u r e s that are p r o b a b l y
modified petioles (leaf s t e m s ) and are k n o w n as p h y l -
lodia. T h e s e usually appear at the end of the g r o w i n g
season and are o f t e n r e f e r r e d to as " w i n t e r l e a v e s , "
since they r e m a i n t h r o u g h o u t d o r m a n c y . T h e p h y l l o -
dia, w h i c h a s s u m e various s h a p e s , can be h e l p f u l in
d e t e r m i n i n g species.
22 / Carnivorous Plants of the United States and Canada
T h e p r o m i n e n t , o f t e n highly decorative trap leaves
are tubular, appearing s o m e w h a t like elongated f u n -
nels or cornucopias. Located at the top is a lobe called
a lid or hood. T h i s is usually supported on a n a r r o w e r
column of v a r y i n g p r o m i n e n c e . T h e hood or lid may be
reflected over the pitcher opening or m a y b e , as it is
in one species, vertical. T h e lid is immobile. R u n n i n g
d o w n the axial seam (that which faces the center of
the plant) of the pitcher leaf is another lobe which is
flattened and w i n g l i k e , its size and character varying
w i t h the species and growing conditions. T h i s struc-
ture is k n o w n as an ala or wing. Pitchers m a y be
colored from green to shades of red, yellow, or
w h i t e . T h e pitchers a s s u m e t w o general habits ac-
cording to s p e c i e s : either erect or nearly erect to total-
ly d e c u m b e n t .
T h e pitchers possess several interesting adaptations
w h i c h serve to lure and entrap prey. T h e bright colora-
tion of the p i t c h e r and the secretions of nectar along
the m a r g i n s of the h o o d , the rolled lip opposite the
c o l u m n , and in some cases the free margin of the ala
lure ground and flying prey to the pitcher opening. T h e
inside of the lid is lined by stiff, downward-directed
h a i r s , w h i c h encourage descent and discourage ascent.
T h e s e hairs vary in size and effectiveness according to
the species. C o n t r a r y to c o m m o n belief, the lid does not
snap d o w n to close off the pitcher after the insect is
c a u g h t . Its exact f u n c t i o n is not totally clear, but it
does, to a degree, p r e v e n t the contents of the pitcher
f r o m being diluted by rainwater in species where the
Fig. 3-1. Sarracenia oreophiia. An example of a pitcher
plant with an erect habit.

Fig. 3-2. S. purpurea ssp. venosa. A common pitcher plant

showing a decumbent habit.

Fig. 3-3. S. purpurea ssp. venosa. Notice the hood with

dense, long, stiff hairs lining the inside and pointing down-
ward.

Fig. 3-4. Longitudinal section of pitcher of S. purpurea ssp.

venosa, showing four basic zones: the inner surface of the
hood lined with stiff, downward-pointing hairs; a smooth
waxy zone; a glandular and absorptive zone without cuticle;
and a lower zone of intermeshing hairs.

Fig. 3-5. Comparison of phyllodia {^'winter leaves").

Smaller, sharply curved S. oreophiia at top; longer and
straight S. flava below.
lid is actually reflected over the pitcher opening. T h e
lid m u s t have a different f u n c t i o n in 5. purpurea, w h e r e
it is vertical.
A f t e r having v e n t u r e d to the b r i m of the pitcher
m o u t h or the underside of the lid, the insect f r e q u e n t l y
o v e r e x t e n d s its f o o t i n g and falls in. T h e interior of the
upper o n e - t h i r d to o n e - h a l f of the pitcher is lined with
a s m o o t h plant w a x w h i c h impedes footing in m o s t
c a s e s , and there is seldom e n o u g h r o o m to begin flight.
D e e p e r in the p i t c h e r , w a x y cuticle is a b s e n t , and the
u n w a x e d s u r f a c e s are capable of a b s o r b i n g digestive
products. All species have a deeper, d o w n w a r d - d i r e c t e d
intermesh of hairs w h i c h f u r t h e r helps prevent egress.
T h e r e are digestive glands that m a y secrete p r o t e i n -
digesting e n z y m e s and fluids of various kinds and c o n -
c e n t r a t i o n s . T h o s e pitchers that d o secrete e n z y m e s
h a v e a small q u a n t i t y of t h e m in their fluids b e f o r e en-
t r a p m e n t o c c u r s , and the c o n c e n t r a t i o n o f e n z y m e s
tends to increase a f t e r initial digestion and a b s o r p t i o n .
T h e relative digestive efficiency of these fluids and e n -
z y m e s f r o m the plant and their possible concert with
microbial activity is still being assessed. ( S e e C h a p t e r
1 , pp. 2 - 3 . )
T h e effectiveness of the trap varies according to
h a b i t u s and species. All the erect traps are e x t r e m e l y
efficient, o f t e n filling to the point that excess insects
can freely w a l k or fly in or out. T h e widely flaring and
reclining pitcher of S. purpurea seems less e f f e c t i v e ;
in f a c t , it is theorized that this species actually drowns
its victims.
T h e digestive m i x t u r e s o f the pitcher are not u n i -
versally effective. V a r i o u s protozoa and insect larvae,
f o r e x a m p l e , have adaptations to resist digestion, and
in fact they b r e e d in the pitcher. A c t u a l l y , the c o n t e n t s
of the pitcher c o m p r i s e s a complex little e c o s y s t e m of
algae, f u n g i , b a c t e r i a , p r o t o z o a , o t h e r m i c r o b e s , and
various resistant insect larvae.
2 4 / Carnivorous Plants of the United States and Canada
T h e flower structure of Sarracenia is generally the
s a m e f o r all species, the only variation being in size,
odor, petal color, and some details of petal shape. T h e
flowers appear in early spring, usually b e f o r e or as
n e w pitcher g r o w t h begins. T h e scapes are tall, to 70
cm in some species, and they support a single nodding
flower. T h e unique floral structure serves to encourage
cross-pollination, although experiments have s h o w n
that artificial self-pollination is quite successful in p r o -
ducing viable seeds. As the spherical flower bud a p -
proaches opening, the scape assumes the shape of a
shepherd's h o o k , and the actinomorphic flower opens
f a c i n g down. T h e r e is an unusual modification of the
style in that the distal half is expanded, so that the
w h o l e looks like an o p e n e d , inverted umbrella and is
c o m m o n l y referred to as such. T h e umbrella has five
points b e t w e e n w h i c h hang the pendulous, elongate
p e t a l s , w h i c h are strap-shaped to o b o v a t e (rounded,
b u t wider than l o n g ) . At each umbrella point is a
small, V - s h a p e d cleft, at the lower point of which is
located one very small stigmatic lobe. T h e rounded
ovary at the base of the style and the numerous sta-
m e n s are located inside a sort of floral c o m p a r t m e n t ,
at the top of which are the five sepals, three b r a c t s , and
the b a s e s of the p e t a l s ; the pendulous portions of the
petals h a n g along the sides of the c o m p a r t m e n t like
drapes, and b e n e a t h is the cupped, expanded umbrella.
At the b a s e s of the petals are nectar-secreting glands.
T h e insect pollinator, o f t e n a b e e , is p r o b a b l y at-
tracted by the color of the petals, the nectar, and the
odor, and usually lands at the bases of the petals b e -
n e a t h the frequently reflexed sepals, where it circum-
navigates the flower several times. It finally enters
at the only visible n a r r o w parting of the petal " c u r -
t a i n s , " over a point of the umbrella, and thereby
b r u s h e s a n y pollen collected from previous flowers
onto the stigma lobe. Inside the flower, pollen has b e e n
shed from the overhanging stamens and has fallen to
the floor of the cupped umbrella, where the bee m a y
collect it, or it m a y collect it while investigating the
stamens and nectar glands near the flower base. W h e n
the bee leaves the flower, it m a y not retrace its steps
by passing over one of the umbrella tips and stigma
lobes, since this offers poor footing prior to flight. In-
stead, the insect exits more often by pushing out one
of the pendulous petals and flying from the wider
umbrella edge between t w o points, thereby mini-
mizing the chance of depositing newly collected pol-
len on the plant's own stigma.
A f t e r pollination and fertilization of the flower,
the petals drop, and the flower frequently assumes a
more nearly or totally erect position. T h e sepals and
umbrella remain all season as the seed matures. Even
if the flower has not been fertilized, the petals fall
within two weeks of the flower's opening. In autumn,
the brown, tubercular seedpods split at five seams and
shed teardrop-shaped, 2 m m , light brown to pink-gray
seeds. Dispersion is probably most often by gravity
and water. T h e seeds require a period of stratification
(damp winter chilling) before they will germinate. This
characteristic effectively prevents the freezing of ten-
der seedlings during the winter months.

G E N E R A L . — A bog or savannah of tall, golden-yellow

S. flava or brilliant, white-topped S. leucophylla g r o w -
ing so thickly that one cannot walk without stepping
over plants, is indeed an attractive and often startling

Fig. 3-6. Flower of Sarracenia flava, typical of the genus.

One petal has been removed to disclose the umbrella, the
stigma point, and the stamens.

Fig. 3-7. Umbrella of S. flava flower spread out to show

points and stigma lobes.
sight. Equally interesting, on a " t r o t " through a n o r t h -
ern sphagnum bog, is coming upon clumps of gaping
pitcher mouths of S. purpurea, which are sometimes
just above sphagnum level while the rest of the plant
is nearly buried in live red-and-green moss. Pitcher
plants are certainly our largest and among our most
easily observed carnivorous plants, but unfortunately
they are very susceptible to change in their habitat.
Very few of the heavily populated, multi-acre stands
described earlier in the century are still extant in the
southeastern ranges. M o r e often, one will find only
scattered clusters or individuals, sometimes clutching
the bank of a ditch that has drained a nearby savannah.
Drainage and fire prevention allow dryland plants
to m o v e in, and these then compete with Sarracenia.
T h e natural late summer and fall fires of years past
helped prevent such recolonization by clearing the site
Fig. 3-8. A dense stand of S. flava in Georgia.
Fig. 3-9. A multicrowned plant of S. flava. Note the short
grass kept cropped by overgrazing cattle, which do not eat
the pitcher plants.
of debris and competing plants while the underground
stems of the pitcher plants were protected. M a n y field
experiments have demonstrated the beneficial effects
of regular fires in a bog or savannah. There is some
indication that fire m a y also release minerals tied up in
dry, dead pitcher leaves and dead insects so that rains
m a y leach them into the soil.
Regarding the effects of the control of competitors,
I recall a situation in Georgia where a landowner had
fenced off a section of savannah in which he was graz-
ing cattle. He noted that the " l i l i e s " (S. flava) were
increasing inside his fence where the cows were g r a z -
ing, while outside, the plants were decreasing. His
puzzlement was further compounded by the failure of
a yearly firing of the pasture to control the pitcher
plants. Actually, he was burning competing plants,
and the cattle were devouring all the grasses, herbs,
and young woody plants—in other words, weeding.
His misguided attempts to eliminate the pitcher plants
in order to g r o w a good stand of pasture grass actually
maintained a virtual garden of Sarracenia.
As we indicated, m a n y small animals are capable of
bypassing a pitcher or even converting it to their own
use. Small spiders, snails, slugs, and frogs may visit
the lips of the pitcher openings in search of food or
prey. The occasional report of frog skeletons in pitcher
contents indicates that circumvention is not always
successful.
The larva of a fly (appropriately named Sarcopha-
ga) resists digestive action by the secretion of anti-
enzymes. It feeds on the debris and pupates in the
depths of the pitchers, doing no h a r m at all. People
tell of opening " l i l i e s " to remove the m a g g o t and using
it as fishing bait.
Within some pitchers, an unusual grass-cutting wasp
builds a condominium of incubators consisting of al-
ternate layers of dry grass and egg c o m p a r t m e n t s
with paralyzed crickets for the larvae to feed upon
when they hatch.
But the greatest real attack on pitchers comes f r o m
a small yellow-and-black moth, Exyra. T h e r e are three
species, each attacking different groups of species of
Sarracenia in different ranges. The adult m o t h is able
to walk about freely on the slippery, w a x y surface of
the inner pitcher where it hides during the day. If dis-
Fig. 3-10. Pitcher of S. flava opened to show infestation by
turbed, it will back down the pitcher still further. If unusual, grass-cutting wasp Isodontia. Plugs of grass alter-
removed, it will promptly flutter to another pitcher and nate with stunned insects for the larvae to feed on when
seek a new hiding place. The female lays one egg per the eggs hatch.
pitcher, and the larva wreaks havoc.
Fig. 3-11. Pitcher of 5. flava infested with larva of Exyra
The larva often spins a dense web across the pitcher moth. The top has collapsed because the inner layers of
mouth, closing off any further trapping. T h e n it feeds the wall have been consumed.
on the inner layer of pitcher tissue, causing the dried,
papery top to fall over and seal the pitcher off from Before it pupates, the larva cuts two holes in the
rain and interference. T h e pitcher becomes a private lower portion of the pitcher: one above the a c c u m u -
feeding and rearing area. T h e brown, collapsed tops lated waste frass, to be used for escape as an adult
of the pitchers are telltale signs of infestation, and moth (moths do not have cutting mouth p a r t s ) , and a
many bogs are severely afflicted. In very severe cli- lower hole for drainage in case some water seeps in
mates, the larva may leave a mature pitcher and winter during the wet winter season. But there is partial re-
over for several seasons in the dead remains of previous buttal by natural balances. Some birds have learned
y e a r s ' growth. In the spring, it enters at the apex of a that the holes mean larval or pupal food within, and
newly developing pitcher before it opens and girdles slash marks made by beaks indicate that m a n y of
the top. these parasitic insects never reached adulthood.

27 / The Eastern North American Pitcher Plants

 Since m a n y larvae of Exyra do overwinter in the D E S C R I P T I O N . — T h e pitchers are curved and decum-
fallen, brown pitchers of the previous year, fire is bent, measure to 45 cm, and widen very prominently
again helpful to the plant colony by burning old litter toward the mouth. There is a large, often slightly un-
from the previous growing season and destroying the dulate ala. T h e hood rises vertically and is lined by
harmful insects. Indeed, the m o s t infested stands of long, stiff hairs. T h e edges of the hood are quite un-
plants are frequently those protected f r o m fire. dulate and have lateral wings. T h e color of the pitchers
T h e whole field of associations between pitcher varies f r o m bright yellow-green to dark purple and is
plants and all the components of their environment is most commonly a middle variation with strong red
a fascinating study with a great deal yet to be explored. venation.
I have never gone into the field without returning with Flower petals, sepals, and flower bracts are mainly
some new perspective, question, or idea. rose pink to dark red. (See exceptions below.) The
flower has a moderate odor of mixed nature, both feline
or musty on one hand, and sweet on the other. Some
The Species
have compared it to that of green peaches or peach
twigs. T h e sweet component is most easily detected
Sarracenia purpurea L. early in the morning or in shaded plants, while the
BOTANICAL NAMES: Sarracenia purpurea ssp. pur- feline odor becomes predominant in bright sun or as
purea Wherry; Sarracenia purpurea ssp. pupurea f. the day progresses.
heterophylla (Eaton) Fernald; Sarracenia purpurea ssp.
venosa Raf. These are three more or less recognized
entities within the basic species.
C O M M O N N A M E S : N o r t h e r n pitcher plant, southern
pitcher plant, sidesaddle plant, pitcher plant, h u n t s -
man's c a p , frog's britches, d u m b w a t c h e s . (The last is
an interesting local term used in the N e w Jersey Pine
Barrens, where the expanded style and sepals that re-
main after flowering were thought to look like open
watches without hands, and therefore mute.)
R A N G E : T h e species has a wide distribution as far
west as northeastern British Columbia, where it was
recently found, and over a good part of the eastern
third of the United States and C a n a d a , with an u n e x -
plained skip area in middle eastern Georgia.
F L O W E R I N G S E A S O N : F r o m as early as M a r c h in
its far southern range to July or August in the north.
T R A P S E A S O N : T r a p s tend to be evergreen unless
unduly exposed.

28 / Carnivorous Plants of the United States and Canada

 Since the species covers such a wide territory, it is S. purpurea ssp. purpurea has narrower, longer
logical that there should be some evolutionary varia- pitchers, which are smooth and glabrous (hairless) on
tion on the periphery of the range. T h e r e is, and the the exterior. T h e diameter of the hood when the wings
variations have been variously interpreted as separate are pinched together does not exceed the diameter of the
species, subspecies, forms, or of no real significance. pitcher. Venation is frequently present when plants
The preponderance of evidence leads me to the follow- are growing in full sunlight, and there m a y be a diffuse,
ing interpretations for the present: T h e plants of the coppery purple color to the upper pitcher. W i n t e r i n g
northern extremity of the range should be designated pitchers of the n o r t h e r n subspecies often turn deep
S. purpurea ssp. purpurea, a noteworthy form being S. m a r o o n and become green again in the spring.
purpurea ssp. purpurea f. heterophylla; the southern T h e form of the northern subspecies, S. purpurea ssp.
plant should be designated S. purpurea ssp. venosa. purpurea f. heterophylla, is found in one county in

Fig. 3-12. Two pitchers of S. purpurea ssp. venosa growing

on the surface of a pond in the New Jersey Pine Barrens.

Fig. 3-13. Flower of S. purpurea, typical of a red-flowered

Sarracenia.

2 9 / The Eastern North American Pitcher Plants

Michigan and in some eastern Canadian bogs in lim-
ited numbers, even though it m a y tend to dominate
in an individual bog. This plant is without any red
pigment at all, the pitchers and flower parts being yel-
low to yellow-green. Intermediates or form hybrids
are easily found in such bogs if the typical subspecies
is also present.
The southern plant is designated S. purpurea ssp.
venosa. T h e pitchers are wider and stockier and have
more prominent, coarse, red venation, always a fine
coat of wooly hair (more easily felt than seen) on the
exterior, and much more expanded hood wings.
This seems to be the best classification consistent
with a broad perspective of our present knowledge.
The two major subspecies are widely separated at the
extremities of their ranges, and they have important
biological differences and distinctive adaptive features
that cannot be ignored.
W h e n plants of one extremity of the range are trans-
planted to the other, they generally retain their char-
acteristic features, but in a relative w a y ; there is a
tendency for the transplant to come to resemble super-
ficially the native. Therefore, local soil, water, and cli-
matic conditions do play some part in determining
the plants' forms, but always within the governing
genetic framework of any group of plants.
W h e r e the ranges of the two subspecies merge in
the N e w Jersey Pine Barrens, one can find both forms
either in separate bogs or often side by side in the same
bog. There are, of course, m a n y interbred intermediates
as well.
Parallel to these observations, the species is host
to the larva of a harmless mosquito of the genus Wye-
omyia, one species of mosquito reportedly inhabiting
the pitcher fluid of the northern plant, and another
species, that of the southern. It has been suggested
that, where the two subspecies intermingle in the N e w
Fig. 3-14. Comparison of pitchers of S. purpurea ssp. venosa
on the left, and ssp. purpurea on the right. Note that the
latter is generally longer and narrower and has a smaller
mouth and less expanded hood wings.
Fig. 3-15. S. purpurea ssp. purpurea /. heterophylla growing
in an open Michigan bog. Note complete lack of red
pigment.
Jersey area, each species of mosquito is able to select Sarracenia flava L.
and stay with its appropriate plant. H o w e v e r , these
BOTANICAL NAME: Sarracenia flava L.
initial reports have recently been questioned.
C O M M O N N A M E S : Yellow trumpet, trumpet, h u n t s -
m a n ' s h o r n , lily.
GENERAL.—Looking at the pitcher of 5. purpurea,
R A N G E : T h e species is generally confined to the s o u t h -
one would guess that, of all the pitcher plants, this
eastern United States in an arc of the coastal plain
one would be the least efficient. It apparently has the
f r o m Virginia through the Florida panhandle into the
weakest enzyme secretions and depends heavily on
Mobile B a y area, but it can be found in some relic bogs
bacterial action for the digestion of its prey, or so
in the southeastern piedmont. It has been planted in,
present evidence seems to indicate. It traps its prey
and has adapted to, outdoor bogs in Pennsylvania.
by drowning it. Since the mouth of the pitcher is widely
T h e r e are claims of adaptation even farther north, but
exposed to weather and flood, rainwater can easily
the y e a r - t o - y e a r persistence and quality of the plant-
dilute or overflow the contents of the pitcher. But these
ings is disputed.
are relatively synthetic observations which are p r o b -
ably not pertinent in the end. W h e n one sees large
populations in good strong bogs and notes the wide
area of distribution, one cannot help agreeing that in
spite of its clumsy appearance the species has certainly
adapted and flourished. We have to learn more about
its adaptations.
The most luxuriant stands of S. purpurea are in the
northern reaches, where there are frequently massive
clusters of plants with multiple crowns a meter or more
in diameter. The species is also seen growing in dense,
floating mats on water at the edges of bog ponds and
lakes and nearly all the w a y across slower, acid
streams.
In the southeast, there is an incompletely studied
race of the species with large, diffusely red to purple
pitchers in sun or even in shade. T h e flower petals are
a pale pink and tend to be sharply curved around the
outside of the pale green umbrella, rather than pen-
dulous. F L O W E R I N G S E A S O N : M i d - M a r c h in the south to
T h e species is often found adapted to alkaline marl mid-April and M a y in the Carolinas.
bogs around the Great Lakes, where the pitchers are T R A P S E A S O N : T h e pitchers brown and wither a t
more numerous, smaller, more brightly colored, and frost. Straight phyllodia are produced in late summer
brittle. and persist all winter.

31 / The Eastern North American Pitcher Plants

DESCRIPTION.—S. flava has an erect, handsome
pitcher which g r o w s up to 90 cm. T h e pitcher has a
wide, flaring m o u t h and a broad, nearly flat, well-
formed lid with a prominent keel and a strong column
with backward-reflexed margins. The ala is much re-
duced. This species has considerable polymorphism
(vein and color variation), particularly in the Carolinas,
with less elsewhere in its range.
T h e r e are four typical forms. The most c o m m o n is
pale green to bright yellow when growing in full sun,
and there is a large m a r o o n splotch on the inside of
the column, from which red veins radiate locally. T h e
next most c o m m o n form has a bright to deep red color
on the external surface of the lid and column, s o m e -
times extending down the pitcher to the ground. V e n a -
tion is moderate, and the m a r o o n color spot is weak. A
third variant is uniformly golden-yellow in full sun-
light, with such coarse and prominent red venation all
over that the surface has a pleated or reticulate a p -
pearance. Again, the interior column spot is weak.
Finally, there are plants with no red pigment at all, the
mature pitchers being uniformly pale green to yellow.
All these basic forms can occur in the same stand, al-
though one or the other frequently predominates, and
hybrid intermediates are easily seen. These variants
have not been clearly named and are under further
study.
The flower is large, and the petals are strap-shaped
and bright yellow. There is a very strong feline odor
which is noticeable from some distance. The plant p r o -
duces ensiform (straight) phyllodia in late summer,
and these persist through the winter. This fact has not
been generally noted in the literature.
G E N E R A L . — T h e r e are areas in Georgia where one
can still see magnificent stands of the species—tall,
bold, bright yellow pitchers filling a large savannah
3 2 / Carnivorous Plants of the United States and Canada
and melting to a vast golden blur when viewed from a
distance. S. flava is especially susceptible to the preda-
tions of the m a n y insect larvae previously mentioned,
but deleterious effects on the species as a whole are not
perceptible. Unfortunately, the species does not re-
spond to attacks on its habitat with equal fortitude,
and prime lots of 5. flava are rapidly disappearing, es-
pecially in the Carolinas, which were once a main
stronghold. This point was illustrated not long ago
w h e n an experienced field botanist and I were in what
is left of the Green S w a m p of eastern N o r t h Carolina.
We were to finish our walks that day by visiting a fa-
vorite location of his for Sarracenia. After parking the
car and hopping the inevitable drainage ditch, we
scoured the savannah for half an hour or so and did
not see one pitcher plant of any species. He was stand-
ing with a look of bewilderment on his face, and when
I came over to him he shook his head and commented,
" W h e r e ' v e they all gone? Y o u could always come here
when y o u wanted to see Sarracenias
Fig. 3-16. Yellow, heavily veined form of S. flava. Note that
the purple area of the throat is not diffuse but is a con-
fluence of veins.
Fig. 3-17. Form of S. flava lacking any red pigment.
Fig. 3-18. Intact flower of 5. flava.
Fig. 3-19. Typical form of S. flava. Note the purple pigment
in the "throat" of the pitcher.
Fig. 3-20. Red-topped form of S. flava beside typical plants.
33 / The Eastern North American Pitcher Plants
 Sarracenia alata Wood In good sunlight, the pitcher has a pale yellow-green
color, is finely red-veined, and the inner lid and column
BOTANICAL NAME: Sarracenia alata Wood. Unac-
frequently have diffuse, deep red coloring, as opposed
ceptable synonym: Sarracenia sledgei.
to the purple splotch in S. flava.
C O M M O N N A M E S : Pale pitcher plant, f l y c a t c h e r .
T h e flower petals are c r e a m y to yellow-white and
R A N G E : This species begins on the Gulf coastal plain,
are obovate, rather than strap-shaped as in S. flava.
where S. flava leaves off in southern A l a b a m a , and
There is a m u s t y odor much like that of S. flava, but it
continues into east T e x a s . T h e r e is a n a r r o w area of
is only about half as strong. There commonly are no
overlap with S. flava just east of Mobile Bay.
winter phyllodia.

G E N E R A L . — S . alata seems quite adaptable and is often

found growing in rather dense clay soil as well as in
sandy savannahs. It can be seen in large, striking
stands on slight slopes and in fields along the high-
w a y s of southern Mississippi. M o s t of these western
plants have the deep red color in the upper, inner
pitchers, and m a n y locations appear literally red with
them.

Fig. 3-21. A clump of Sarracenia alata.

Fig. 3-22. S. alata.

Fig. 3-23. S. alata with diffuse, deep red pigment of inner

F L O W E R I N G S E A S O N : Early M a r c h through April. lid and column.
T R A P S E A S O N : Because o f its e x t r e m e southern lo-
cation, the pitchers of S. alata often remain over winter
in protected areas, withering back with frost in more
open places. Year-old pitchers m a y assume a deep red
coloration.

D E S C R I P T I O N . — T h e pitchers are erect and measure

to 75 cm tall. At first glance, this species can easily be
mistaken by the inexperienced for S. flava, especially
in the Mobile Bay area. H o w e v e r , the pitcher m o u t h
of S, alata does not flare as widely, the column is not as
tall or reflexed, the lid is smaller and more c o n v e x ,
and there is a larger ala (hence the specific name, alata).

3 4 / Carnivorous Plants of the United States and Canada

3 5 / The Eastern North American Pitcher Plants
 Sarracenia oreophiia (Kearney) Wherry The flowers have pale yellow-green petals and a
very weak odor easily detectable (by nonsmokers),
BOTANICAL NAME: Sarracenia oreophiia (Kearney)
which is best described as mixed sweet and musty (as
Wherry.
in S. purpurea), the mustiness tending to predominate.
COMMON NAMES: Flycatcher, green pitcher plant.
T h e r e are phyllodia which appear in midsummer.
R A N G E : T h e species is n a r r o w l y confined to a few
These are sharply curved and unique for the species.
scattered locations in northeastern Alabama. Disjunct
sites in m i d d l e w e s t e r n G e o r g i a w e r e r e p o r t e d b u t are
G E N E R A L . — T h i s species was thought to be a form of
no longer extant.
S. flava until 1 9 3 3 , w h e n E. T. W h e r r y discerned that
it was not and formally described it. Its range is g e o -
graphically quite separate from that of S. flava, and
in fact, no other pitcher plants grow with it except in

Fig. 3-24. Sarracenia oreophiia. Note the fine venation and

rather close resemblance to forms of S. flava, except for
smaller lid and less well-developed column.

FLOWERING SEASON: Mid-April to early June.

TRAP SEASON: The pitchers wither quite early,
usually in midsummer. Curved winter phyllodia are
regularly p r o d u c e d in late s u m m e r .

D E S C R I P T I O N . — T h e pitcher is erect and measures

up to 75 c m , with a widely flaring mouth, a well-
formed column which is not as strongly reflexed as in
S. flava, and a large lid with a prominent keel. T h e lid
tends to slope slightly more u p w a r d than in S. flava.
T h e pitchers are very often pale green, but there are
stands where there is a fine, red venation. The ala is
not prominent.
one very small station near Birmingham, where it Sarracenia minor Walt.
meets t h e r a n g e of 5. rubra. S. oreophiia's montane lo-
BOTANICAL NAME: Sarracenia minor Walt.
c a t i o n d i s j u n c t f r o m t h a t o f S . flava, a n d i t s c o n s i s t e n t
COMMON NAME: Hooded pitcher plant.
morphologic and biologic differences, including the
RANGE: The southeastern coastal plain from the
sharply curved phyllodia and the differences in the
southernmost tip of eastern North Carolina to the
c o l o r a n d o d o r o f t h e f l o w e r s , all s u g g e s t a s e p a r a t e
mid-Florida panhandle. This is the only species ex-
species.
tending into the Florida peninsula.
S . oreophiia h a s s o m e i n t e r e s t i n g a d a p t i v e b i o l o g i c a l
features. It grows in wet depressions along streams of
the elevated Sand Mountain plateau of northeastern
A l a b a m a , w h e r e t h e soil i s s a n d y c l a y . I n m i d s u m m e r ,
so-called dog d a y s befall the area, bringing v e r y hot,
humid weather with decreased rainfall and consider-
able drying. At this point, the pitchers begin to b r o w n
and wither while other species of Sarracenia in more
favorable downland areas are still thriving. The
prominently curved phyllodia appear and remain green
all autumn and winter. T h e plants catch what prey
they can early, then literally fold up for the hot, d r y
weather. This early withering of the pitchers is largely
carried over in cultivation in spite of adequate w a t e r
and favorable temperatures.
O n e i s n o t likely t o c o m e u p o n this species v e r y
casually, since there were a l w a y s relatively few stands
in a very small range, and these are disappearing r a p -
idly as land is c l a i m e d for a g r i c u l t u r e a n d t h e g r o w t h
of timber. S. oreophiia is on the list of very much F L O W E R I N G S E A S O N : Late March to mid-May. This
e n d a n g e r e d species a n d will s o o n d i s a p p e a r f r o m n a t u - is also the only species in which flowering m o s t c o m -
ral sites. F o r t u n a t e l y , it is a d a p t i n g in several p r i v a t e monly occurs simultaneously with or slightly after
and botanical gardens in different parts of the world, pitcher growth.
in collections made from seeds and plants gathered T R A P S E A S O N : P i t c h e r s t e n d t o p e r s i s t all w i n t e r i n
from condemned or fading areas. But a good natural sheltered stands in the southernmost range, but else-
stand of a n y species of plant is preferred to a cultivated w h e r e , t h e y g e n e r a l l y die b a c k d u r i n g s e v e r e w i n t e r s .
one. For these reasons, we were especially distressed
to learn that a commercial nursery h a d possibly m a d e D E S C R I P T I O N . — S. minor has an erect pitcher av-
a massive illegal collection from an Alabama state eraging about 25-30 cm but growing up to 80 cm in
park! certain areas. T h e r e is a prominent ala, and the h o o d

37 / The Eastern North American Pitcher Plants

is extremely reflected over and approximated to the
elliptical mouth of the pitcher. T h e column is barely
discernible, being quite short, nonreflexed, and c o n -
tinuous with the hood and body of the pitcher. T h e
color is generally green with a coppery red cast to the
upper pitcher when it is growing in bright sun. There
is fine, red venation over the interior of the hood and
the column. Older pitchers frequently turn dark red
in late summer. T h e flower is odorless and has pale
yellow-green petals. T h e r e are no phyllodia.
G E N E R A L . — A n o t e w o r t h y characteristic of this spe-
cies (and of two others, 5. psittacina and S. leucophylla,
yet to be discussed) is the presence, on the back of
the upper one-third of the pitcher, of irregular, reticu-
late, clear to whitish areas that lack chlorophyl. These
are called light windows, fenestrations, or areolae. All
sorts of functions for the fenestrations in S. minor
have been suggested, but the most reasonable is that
they admit light into the interior of a pitcher rather
darkened by the close approximation of the well-
developed hood. Insect prey are less likely to fly or
crawl into a darkened area than into a lighted one.
W h e n alighting or crawling to the top of the broad
ala, they m a y mistakenly confuse the light windows
for a place of exit and take off in that direction, strik-
ing the pitcher's back wall and falling in. T h e closely
approximated hood, while darkening the interior, u n -
doubtedly provides excellent cover against rain enter-
ing the pitcher.
S. minor seems to have a particular affinity for ants.
On w a r m , clear days, one can go to the field and see
single columns of ants in t w o - w a y traffic up and d o w n
the ala, which is studded on its narrow edge with glis-
tening nectar glands. T h e ants travel all the w a y to
the lip, where m a n y fall in. It is not known whether
this represents a specific attraction or whether it is a
38 / Carnivorous Plants of the United States and Canada
result of S. minor's tendency to grow in a drier en-
vironment where ants are more likely to be.
S. minor is frequently found in the more drained,
upland parts of savannahs or in light pine woods,
whereas most other species of Sarracenia are most
abundant in moister, open areas. There is a question
as to whether this is a " p r e f e r e n t i a l " adaptation to allay
competition with other pitcher plants which do poorly
in drier or shadier locations, or whether S. minor's
adaptation to a dry, shady habitat simply coincides
with the maladaptation of the other species. I have
observed and g r o w n S. minor for a number of years
in both wet and dry soils and in sphagnum, and the
plants are m u c h larger and more robust when grown
in wetter substrates. In fact, the plants reach their
zenith in size in the very wet habitat of the Okefenokee
S w a m p , where they g r o w intermixed with S. psittacina
on huge floating sphagnum islands called prairies.
These Okefenokee plants are a special case, and test
plants have not been removed and studied in con-
trolled transplant experiments to see if their marked
difference in size is really environmental or genetic. I
can speak with more assurance with respect to the
savannah plants of the rest of the coastal plain, with
which I have done transplant studies, and conclude
that these plants are hardier in wet substrates, es-
pecially in sphagnum moss tubs and plantings, and that
therefore the drier location in which S. minor is often
found is probably not a preferential adaptation.
Although its range overlaps that of many other spe-
cies of Sarracenia, there are places, particularly in
South Carolina and parts of the Florida peninsula,
where fine, almost pure stands of S. minor can yet be
seen.
Fig. 3-25. Sarracenia minor, with flowers in early spring. Fig. 3-26. S. minor. Note the well-developed hood closely
approximated to the mouth, and the light windows.

39 / The Eastern North American Pitcher Plants

 Sarracenia psittacina Mich. t h a t line t h e i n t e r i o r o f t h e p i t c h e r a r e t h e m o s t d e v e l -
o p e d i n a n y s p e c i e s o f Sarracenia. E x t e r n a l l y , t h e c o l o r -
BOTANICAL NAME: Sarracenia psittacina Mich.
ation is green in the shade to deep red in good sunlight,
C O M M O N N A M E : Parrot pitcher plant.
and light w i n d o w s are prominent. (See the preceding
RANGE: An arc of the southeastern coastal plain
s e c t i o n on S. minor f o r a d i s c u s s i o n o f l i g h t w i n d o w s
from Georgia through the western Florida panhandle
and their possible function.)
into southern Mississippi.
T h e flower is rather small, with petals deep to bright
red, a n d there is a w e a k but definitely pleasant sweet
odor. There are no phyllodia.

GENERAL.—S. psittacina h a s a very distinctive pitch-

er, its e x t e r n a l a p p e a r a n c e b e i n g r a t h e r unlike t h a t o f
a n y other m e m b e r of the genus except in individual
m o r p h o l o g i c p o i n t s . A side v i e w of the pitcher discloses
a case for the c o m m o n n a m e , parrot pitcher plant. I
h a v e seen the largest, m o s t globose hoods on plants
that have been m o v e d to a piedmont Carolina bog,
even though the natural range is far to the south.

Fig. 3-27. Sarracenia psittacina. Note globose hoods.

F L O W E R I N G S E A S O N : Late M a r c h t o M a y .
T R A P S E A S O N : T h e pitchers remain all year.

D E S C R I P T I O N . — T h e pitchers are d e c u m b e n t ( e x -
cept in extreme shading) and grow to 20 cm, al-
t h o u g h specimens with 3 0 c m pitchers can occasionally
be found. T h e r e is a very prominent, undulate ala. T h e
h o o d is the m o s t elaborate in any species of Sarracenia,
with the edges united so t h a t it is a globose s t r u c t u r e
with a relatively small aperture at the top of the ala.
Inside, t h e a p e r t u r e is surrounded by a 0 . 5 cm collar,
which enables the trap to w o r k s o m e w h a t like a lobster
pot. T h e d o w n w a r d - p o i n t i n g , intermeshing, stiff hairs

4 0 / Carnivorous Plants of the United States and Canada

 This species prefers a very wet habitat, and the pros-
trate rosettes are often flooded in the spring. This seems
to inconvenience the plant little since m a n y aquatic
animals are found trapped inside the pitchers. Some
botanists feel that flooding is a definite advantage, if
not a requirement, for the species. In this situation, one
can speculate how the especially large intermeshing
interior hairs and the collar surrounding the interior
of the aperture are aids in preventing swimming ani-
mals from retreating.
Fig. 3-28. S. psittacina. Less intensely colored, shade-grown
plants.
Fig. 3-29. Longitudinal section of pitcher of S. psittacina.
Note the especially well-developed intermeshing trap hairs
inside.
As in the case of S. minor, this species grows to
rather large size on the floating sphagnum prairies of
the Okefenokee S w a m p , where the two species occur
sympatrically. V e r y large specimens have also recently
been found in bogs in southern Mississippi. Again,
the question of whether the size factor is environ-
mental or genetic will have to be worked out.
The plant is rather abundant within its range, but
it is easily overlooked since it is low on the ground
and, by summer, often deep in grass and other plants.
T h e taller flower scapes are a definite help in locating it.
 Sarracenia leucophylla Raf. nent except in pitchers that appear very early in the
spring and in s h a d e - g r o w n specimens. T h e most strik-
BOTANICAL NAME: Sarracenia leucophylla Raf.
ing characteristic is the white coloration of the hood,
Unacceptable synonym: Sarracenia drummondii.
column, and upper regions of the pitcher; hence the
COMMON NAME: White-topped pitcher plant.
common as well as the specific name, leucophylla,
R A N G E : T h e s o u t h e a s t e r n coastal plain f r o m s o u t h -
m e a n i n g w h i t e leaf. T h e r e are variations ranging f r o m
western Georgia into the w e s t e r n Florida panhandle
upper pitchers that are almost pure white with very dis-
and f r o m the Mobile B a y area to just over the Missis-
sippi line.
Fig. 3-30. A clump of S. l e u c o p h y l l a .

FLOWERING SEASON: Early March to late April.

T R A P S E A S O N : The traps tend to wither in winter,
with n e w traps appearing in two crops, one concomi-
tant with spring flowering and another, stronger set
in late s u m m e r . Phyllodia-like s t r u c t u r e s are o f t e n p r o -
duced during dry periods in midsummer, but they rare-
ly persist over winter.

DESCRIPTION.—S. leucophylla h a s a tall, erect pitch-

e r w h i c h g r o w s t o 9 5 c m . T h e ala i s n o t v e r y p r o m i -

4 2 / Carnivorous Plants of the United States and Canada

crete pale green veins dividing the white area into a G E N E R A L . — T h e r e is no doubt that, with the p o s -
mosaic pattern, to pitchers w i t h r a t h e r h e a v y red veins s i b l e e x c e p t i o n o f b r i g h t y e l l o w S . flava i n n o w r a r e
and m u c h red pigment suffused with purple a r o u n d the m a s s i v e s t a n d s , S. leucophylla i s t h e m o s t e y e - c a t c h i n g
white patches. T h e pitcher m o u t h is quite expanded, o f a n y s p e c i e s o f Sarracenia. Its r a n g e is r a t h e r small,
a l t h o u g h less so in t h e v a r i a n t s of m o r e r e d d i s h color. b u t t h e p l a n t s a r e still a b u n d a n t a n d v e r y f r e q u e n t l y
T h e c o l u m n is well f o r m e d and m o d e r a t e l y reflexed, m a s s e d in large b o g s easily observed f r o m m a n y roads.
a n d t h e lid i s l a r g e a n d w i d e , w i t h a p r o m i n e n t l y u n d u - Where I grow a n u m b e r of species o f Sarracenia to-
late m a r g i n a n d large hairs on the l o w e r surface. gether o u t d o o r s in the piedmont area of N o r t h C a r o -
T h e large flower has deep red petals and a sweet l i n a , I h a v e f o u n d t h a t S . leucophylla i s b y f a r t h e m o s t
odor. attractive to insects. After being open only a few days,
t h e p i t c h e r s b e c o m e n e a r l y full t o the t o p . T h e s a m e

Fig. 3-31. Two color forms of S. leucophylla: white-topped observation applies t o field s p e c i m e n s of S. leucophyl-
with large mouth and green veins, and a smaller form with la, w h i c h g e n e r a l l y c o n t a i n m o r e a c c u m u l a t e d i n s e c t s
red veins. than do the pitchers of other species. T h e curious pat-

Fig. 3-32. A savannah with moderate growth of Sarracenia tern of t w o crops of pitchers a y e a r is also distinctive.
leucophylla, its white tops easily visible above the grass.

43 / The Eastern North American Pitcher Plants

 Sarracenia rubra Walt. T h e p i t c h e r s a r e e r e c t a n d g r o w u p t o 5 0 c m tall
w h e n fully d e v e l o p e d in ideal habitats. M a t u r e pitch-
BOTANICAL NAME: Sarracenia rubra Walt. Also, a
e r s g e n e r a l l y h a v e b r i g h t r e d , fine v e i n s o n a c o p p e r y
generally acceptable subspecies: Sarracenia rubra ssp.
tan background; in less mature specimens the red
jonesii Wherry.
tends to be m o r e diffuse. T h e m a t u r e s u m m e r pitch-
C O M M O N N A M E S : Sweet pitcher plant, red pitcher
ers h a v e a m o d e r a t e l y p r o m i n e n t ala. In the typical
plant.
coastal plain f o r m , the c o l u m n is short and nonreflexed
RANGE: In an arc throughout the coastal plain of
w h i l e t h e lid i s e l o n g a t e , w e l l d e v e l o p e d , a n d s o m e w h a t
the southeastern United States, except the Florida
closely a p p r o x i m a t e d to the elliptical pitcher m o u t h ,
peninsula, and into southwestern A l a b a m a . T h e dis-
a l t h o u g h n o t n e a r l y a s c l o s e l y a s i n S . minor.
junct subspecies is in the southwestern mountain coun-
In the Carolina m o u n t a i n s , w h e r e the controversial
ties of North Carolina and one adjacent county in
jonesii subspecies is found, the pitchers grow 15-20
South Carolina.
cm taller t h a n in the coastal plain f o r m . T h e y h a v e
better-developed, wider hoods, taller columns with
s o m e reflexion, a n d a distinct bulbous widening of the
upper quarter of the pitchers, which can be seen w h e n
the pitchers are v i e w e d f r o m the side.
In the w e s t e r n G u l f coastal plain area, there is a
v a r i a n t f o r m , w h i c h i s a s tall a s t h e m o u n t a i n s p e c i -
m e n s but is a m o r e diffuse red in sunlight a n d h a s a
hood which is better developed, more elongate and con-
vex, and often s o m e w h a t undulate at the margins. The
column is slightly reflexed and better developed than
in the eastern plants, but not as strongly developed as
i n t h e m o u n t a i n jonesii.
In nearly disjunct locations around Montgomery,
Alabama, and extending down to northern Baldwin
F L O W E R I N G S E A S O N : April to M a y in the coastal C o u n t y , there is an additional variation, measuring to
plain, to June in the Carolina mountains. 30 cm, with stockier pitchers and wider openings.
T R A P S E A S O N : T h e pitchers wither with frost. T h e r e i s o n l y a f a i n t r e d c o l o r a t i o n o f t h e lid a n d v e r y
fine v e n a t i o n o f t h e u p p e r p i t c h e r w h e n t h e p l a n t i s
D E S C R I P T I O N . — T h i s is a difficult s p e c i e s to d i s c u s s . g r o w n i n full s u n l i g h t .
There is much animated controversy over the exact R a t h e r r e g u l a r l y , p l a n t s of this entire c o m p l e x s p e -
t a x o n o m i c status of the populations, w h i c h s o m e b o t - cies h a v e a t w o - s t a g e leaf sequence. Spring leaves m a y
anists w o u l d classify a s a c o m p l e x o f species. W e will be somewhat phyllodiform, with a broad ala and a
try to present a practical s u m m a r y of the situation. small pitcher tube. T h e leaf is often d e f o r m e d , being

44 / Carnivorous Plants of the United States and Canada

Fig. 3-33. Sarracenia rubra (typical form of eastern coastal
plain) in flower in early spring. Note the dead pitchers from
previous years at the left, and younger pitchers emerging.

Fig. 3-34. S. rubra, same pond edge as above, later in sea-

son. Petals have fallen from the flowers, leaving the um-
brella and sepals. The pitchers are now better developed
and well colored.

Fig. 3-35. A form of S. rubra peculiar to the Gulf coastal

plain.

Fig. 3-36. S. rubra ssp. jonesii. Afore the more robust pitcher
tops and the definite bulge or widening just below the
mouth.
ensiform, or roughly S-shaped. The summer leaves, a n d S . leucophylla, a r e j u s t a s s t r i k i n g a n d p r o b a b l y a s
which appear later, are m o r e typical of the particular i m p o r t a n t a s t h o s e i n S . rubra, y e t t h e y h a v e r e c e i v e d
form. This dimorphic pattern varies considerably little o r n o a t t e n t i o n . )
among different populations and forms because of In 1929, E. T. W h e r r y inadvertently sparked this
genetic differences, local growing conditions, or a c o n t r o v e r s y w h e n h e p u b l i s h e d his d e s c r i p t i o n o f the
combination of these factors. Reciprocal transplant- new species Sarracenia jonesii for disjunct populations
ing of individuals may elucidate the mechanism in- o f S . rubra i n t h e C a r o l i n a m o u n t a i n s . H e felt t h a t g e o -
volved. g r a p h i c a l s e p a r a t i o n f r o m t h e r e s t o f t h e rubra s t a n d s ,
T h e flowers are rather small and h a v e bright red along with the morphological differences and his in-
p e t a l s . W h i l e m o s t s p e c i e s o f Sarracenia h a v e o n l y o n e correct observation that the flower of the mountain
f l o w e r p e r g r o w t h c r o w n , S . rubra f r e q u e n t l y h a s m u l - p o p u l a t i o n h a d n o o d o r , w e r e all s u f f i c i e n t t o s u g g e s t
t i p l e f l o w e r s i n all i t s f o r m s . T h e p l e a s a n t l y s w e e t o d o r a n e w species, s o m e w h a t parallel to his separation of
is strong except in the Gulf coastal variants, which have S. oreophiia a few years later. Unfortunately, he ap-
only a w e a k odor. T h e r e are no true winter phyllodia. parently m a d e the mistake of believing that the m o r e
r o b u s t G u l f c o a s t a l v a r i a n t s o f rubra a n d t h e d i s j u n c t
G E N E R A L . — P a r t of the problem with the t a x o n o m y central A l a b a m a p o p u l a t i o n s * w e r e identical with the
of this a t t r a c t i v e little p l a n t c o u l d be put into per- mountain plants, which they are not. He has since
spective if m o r e consideration were given to the cur- rescinded his opinion, but S. jonesii is recorded in
rent tendency to consider species as dynamic plant botanical literature as being in the Gulf coastal plain.
populations. Plants are always interacting with each The S. jonesii designation of the Carolina mountain
other and with their e n v i r o n m e n t and are fully in- plant m e t with considerable resistance in the forties,
v o l v e d i n e v o l u t i o n a n d its c h a n g e s ; t h e y a r e n o t the and recent monographs on Sarracenia have since
static entities that symbolic names suggest. An ex- tended to relegate the m o u n t a i n plants to either s u b -
cessive desire for certainty and a need to label c a n s p e c i f i c s t a t u s o r n o s t a t u s a t all.
lead to quibbling which suppresses truly important The confusion was further compounded when the
questions. In most cases, plants do not change or typical S. rubra of the eastern coastal plain was re-
evolve m u c h in one m a n ' s lifetime, or even in m a n y p o r t e d i n m o u n t a i n b o g s a l o n g s i d e jonesii. I a m s a t i s -
l i f e t i m e s , a n d it is difficult to a p p r e c i a t e b r o a d e r c o n - fied t h a t t h e r e a r e n o c o a s t a l p l a i n f o r m s o f S . rubra
cepts than those which are immediately apparent. in these mountain slope bogs, and that there never
S. rubra appears to be exhibiting something of a were. T h e immature pitchers of younger or disturbed
s p r a w l of e v o l u t i o n a r y d i v e r g e n c e a n d p r o b a b l y illus- plants o f jonesii /ook like the coastal ones until the
trates the concept of incomplete differentiation. Per-
h a p s the seemingly equivocal but really quite useful *A recent paper (Case, F. W. and Case, R. B. 1974. Sarracenia
alabamensis, a newly recognized species from central Ala-
terms semispecies and syngameon should be considered
bama. Rhodora 7 6 : 650.) suggests that the separate central
by more students of extreme persuasions as a replace- Alabama plants—also becoming extinct—should be designated
as a species, Sarracenia alabamensis. The authors' studies were
m e n t f o r t h e t e r m subspecies. (Curiously, the variations
not thorough, however, and my impression is that these popu-
of S. flava, and even the lesser ones of S. oreophiia lations will likely prove to be an S. rubra subspecies.

4 6 / Carnivorous Plants of the United States and Canada

plants mature or acclimate. T h e report of both types The whole problem may, before long, become an
and their i n t e r m e d i a t e s w a s o n e o f t h o s e m i s o b s e r v a - exercise in futility. T h e jonesii p l a n t s are nearly all
tions that h a d not been followed up by the n e c e s s a r y extinct. T h e y can be found in only a few small bogs
y e a r s o f c a r e f u l l y o b s e r v i n g m a r k e d field p l a n t s a n d after considerable search where they were o n c e easily
reciprocal transplants. seen in masses f r o m the roadside. Likewise, the typical
Still m o r e r e c e n t l y , t w o t y p e s o f e a s t e r n c o a s t a l p l a i n coastal plain plants are fast disappearing in the east,
f o r m s o f S . rubra h a v e b e e n s u g g e s t e d ! A g a i n , t h i s i s a particularly in the sandhills below the piedmont.
hypothesis based on static h e r b a r i u m studies a n d ir-
r e g u l a r field v i s i t s , r a t h e r t h a n o n l o n g - t e r m o b s e r v a - SARRACENIA HYBRIDS
tions of live, g r o w i n g plants. T h e pitchers simply g r o w T h e m a p s s h o w i n g t h e r a n g e s o f s p e c i e s o f Sarra-
taller a n d a r e m o r e r o b u s t in p l a n t s g r o w i n g in w e t cenia indicate that many occur in the same or over-
s p h a g n u m areas, on pond edges, and in low sandhill l a p p i n g r a n g e s . I n t h e field, o n e will o f t e n n o t e t h a t
bogs than they do in the smaller, almost miniature t w o or m o r e species occur in the same bog or s a v a n n a h .
forms growing in drier savannahs. This has been As a result, crosses or hybrids between two species are
proved to my satisfaction by reciprocal transplants as frequent.
well as by g r o w i n g the plants in o u t d o o r t u b s a n d a N o t all s p e c i e s i n t h e p l a n t k i n g d o m a r e c a p a b l e o f
greenhouse for several years. T h e differences reported hybridizing. W h e n species do cross, the resulting p r o g -
in eastern coastal plain plants are, therefore, non- e n y a r e f r e q u e n t l y s t e r i l e — t h a t is, t h e y a r e i n c a p a b l e
genetic. of further sexual reproduction. Such is not the case
For the time being, I would suggest the following w i t h Sarracenia, h o w e v e r . N o t o n l y c a n n e a r l y all c o n -
guideposts for the beginner and even for m o r e ad- ceivable crosses be found or made in the field and
vanced naturalists w h o are equally confused by the g r e e n h o u s e , but the h y b r i d s are quite fertile a n d are
j i g s a w p i c t u r e in b o t a n i c a l l i t e r a t u r e . T a k i n g 5. rubra capable of forming additional crosses with third spe-
i n p e r s p e c t i v e w i t h all o t h e r m o r p h o l o g i c a l a n d b i o - cies or with each other, or they can effect c o m p l e x
logical considerations in the g e n u s and with relative backcrosses with one or both parent plants. Such back-
differences between accepted species of Sarracenia, crossing (introgressive hybridization) results in an
t h e b a s i c d e s i g n a t i o n S . rubra c a n a p p l y t o all p o p u l a - e x c h a n g e of genetic material b e t w e e n m o r e or less e s -
tions, with a subspecific designation S. rubra ssp. t a b l i s h e d s p e c i e s , a n d i t i s felt b y m a n y b o t a n i s t s t h a t
jonesii f o r t h e C a r o l i n a m o u n t a i n p l a n t s , s i n c e c o n s i s - this is an i m p o r t a n t f a c t o r in creating variation for
tent morphological and biological differences along w i t h evolution.
the factor of isolation c a n n o t be ignored. B u t n o n e of T h e r e is a general rule in b o t a n y that plant hybrids
these are sufficient in t h e m s e l v e s or t o g e t h e r to s u g - b e c o m e e s t a b l i s h e d o n l y w i t h difficulty, since in t h e o -
gest a separate species for jonesii. The Gulf coastal retical ecologic t e r m s (and quite usually in actuality)
variants and the central A l a b a m a plants also d e s e r v e a they should require a rather n a r r o w habitat interme-
subspecific status, a l t h o u g h s o m e b o t a n i s t s are willing diate between those of the two parents, which have
to concede species status, which I believe would be t h e m s e l v e s b e c o m e established as a result of specific
p r e m a t u r e at this point. environmental selection. But the various species of

47 / The Eastern North American Pitcher Plants

Sarracenia can generally grow pretty well in similar o v e r l a p , t h e r e i s r e l a t i v e l y little o p p o r t u n i t y f o r c r o s s -
environments, so the hybrids establish themselves i n g , s i n c e t h e t w o peaks o f f l o w e r i n g a c t i v i t y w i t h i n
intermingled among the parents, provided there are the broad flowering periods are different in m o s t cases.
physical space and proper conditions for seed g e r m i n a - Thus, there would be few plants of sympatric species
tion along with a m i n i m u m of c o m p e t i t i o n f r o m o t h e r able to cross at the critical period. Such an incomplete
species in the bog. T h e h y b r i d s m a y , h o w e v e r , c o m e isolation factor is frequently referred to as " l e a k y . "
to occupy newly opened, disturbed areas. T h e r e are some other equally leaky isolation factors
O n e might ask, then, if these species are so capable that apply t o Sarracenia. A large pollinator capable of
of crossing with each other, h o w distinct species h a v e negotiating a large-flowered species m a y not be able
been preserved so well, especially in s y m p a t r i c b o g s . to enter the flower of a smaller s y m p a t r i c species, al-
T h e a n s w e r lies p a r t i a l l y i n i s o l a t i o n f a c t o r s . I s o l a t i o n though the reverse could occur. Color and nectar odor
most c o m m o n l y involves physical or geographic bar- m a y attract certain pollinators preferentially, thus lim-
riers or displacements. T h e r e c a n also be r e p r o d u c t i v e iting the t r a n s f e r of pollen b e t w e e n species with dif-
isolation, such as a g e n u s ' inability to hybridize, or the ferent flower colors or odors. M a n y pollinators exhibit
f o r m a t i o n o f sterile h y b r i d s o r h y b r i d s t h a t m a y b e a c h a r a c t e r i s t i c k n o w n as fidelity, w h e r e b y a p a r t i c u -
ecologically incompatible with the locality w h e r e their lar colony of pollinating insects will visit only one
seed is shed. B u t we h a v e seen t h a t neither of these species, or even one stand of a species, until it ceases
factors quite applies to s p e c i e s o f Sarracenia occurring to flower, rather than visiting all flowering plants
in the s a m e bog, a l t h o u g h the physical s e p a r a t i o n f a c - indiscriminately.
tor would apply in disjunct bogs of single species and Plant dispersal in Sarracenia in general is also lim-
at the far ends of ranges. i t e d . A l t h o u g h m u c h m o r e field r e s e a r c h n e e d s t o b e
When two or more species o f Sarracenia occur in- done on this subject, the Sarracenia pollinators that
termingled in the same bog, other somewhat more have been studied are not far ranging. Furthermore,
subtle isolation factors c o m e into play. T h e m a i n one the seeds of the genus d e p e n d on floods or n e a r b y
appears to be differences between the peak flowering flowing water to be dispersed. T h e y are too heavy for
periods of the species, which m e a n s that w h e n the pol- wind dispersal, and we do not k n o w of any birds or
len of o n e species is ripe, the s t i g m a of a n o t h e r species m a m m a l s that carry them. O t h e r isolation factors c o m -
m a y not be receptive yet, or the plant m a y not even m o n to the entire g e n u s involve the inability of a h y -
have flowered. Although flowering periods sometimes brid seed crop to b e c o m e established in a c r o w d e d area
w i t h o u t o p e n , d i s t u r b e d sites a n d w i t h o u t s u i t a b l e soil
and water after seed dispersal does occur. Finally,
Fig. 3-37. S. flava x S. purpurea hybrid- m a n y insect larvae attack m a t u r i n g seed pods.

fig. 3-38. S. purpurea x S. rubra. Individually, these isolation factors are seen to be
fraught with loopholes, but if they are taken together
Fig. 3-39. S. minor x S. purpurea.
w i t h o b s e r v a t i o n s i n t h e field, t h e r e i s g o o d e v i d e n c e
Fig. 3-40. 5. leucophylla x S. purpurea x 5. leucophylla. that they are generally effective, even t h o u g h m a n y h y -

Fig. 3-41. 5. leucophylla x S. psittacina. brids are rather easily found.

49 / The Eastern North American Pitcher Plants

 W h i l e inspecting healthy locations of Sarracenia, one vanity) as horticultural names, with the placement of
will notice that certain hybrid c o m b i n a t i o n s occur with an x in the written version, as in S. x catesbaei, the x
variable f r e q u e n c y depending on the application of w a r n i n g that we are not dealing with a species.
any or all of the possible isolation factors m e n t i o n e d In a p p e a r a n c e , Sarracenia hybrids assume a form
a b o v e , and p r o b a b l y on m a n y o t h e r f a c t o r s we do not that is generally i n t e r m e d i a t e b e t w e e n those of the two
yet k n o w about. T h e h y b r i d b e t w e e n S. flava and S. p a r e n t s , rather than representing a patterned mosaic
purpurea m a y be f o u n d with moderate ease in a loca- or dominant-recessive situation. A yellow-flowered
tion where the two species occur together. T h e hybrid species crossing with a red-flowered species does not
b e t w e e n S. flava and S. minor, on the other h a n d , is produce a hybrid with one color or the other, or with
relatively rare where the two species g r o w together. spotted flowers; it produces pink or orange flowers.
S o m e species hybridize so freely where they occur t o - A species with erect pitchers crossing with a decumbent
g e t h e r that so-called h y b r i d s w a r m s are found. T h i s is species produces a hybrid with pitchers that are semi-
frequently the case with S. alata and S. leucophylla, erect.
where several large bogs full of intercrosses and b a c k - T h e r e are large n u m b e r s of natural hybrids that
crosses yield few plants that can be identified as pure have b e e n f o u n d and recorded, and a few possible
p a r e n t s . T h e s e bogs occur at the n a r r o w interface of ones are yet to be f o u n d , but practically all possible
the ranges of the two species, and it is clear that g e o g - c o m b i n a t i o n s have been produced in the laboratory or
r a p h y plays a big role in keeping these two species g r e e n h o u s e . W e have included photographs o f only
intact. a few of the naturally found hybrids as examples.
T h e f r e q u e n c y of hybridization in Sarracenia caused T h e interested reader is invited to consult some of the
a great deal of difficulty a m o n g early b o t a n i s t s , w h o references for more discussion.
o f t e n t h o u g h t that each hybrid was a n e w species and S o m e naturally f o u n d , simple, two-parent Sarracenia
so n a m e d it. O v e r years of s t u d y , the specie* were hybrids and their horticultural n a m e s :
sorted from the h y b r i d s , but some of the original spe-
cies n a m e s given to the h y b r i d s stuck in a modified Botanical n a m e Horticultural name
w a y , and the plants are o f t e n referred to accordingly. S. flava x S. purpurea 5. x catesbaei
For e x a m p l e , a plant n a m e d Sarracenia catesbaei was 5. leucophylla x S. purpurea S. x mitchelliana
later found to be a cross b e t w e e n S. flava and S. pur- S. minor x S. purpurea S. x swaniana
purea. T h e h y b r i d should m o s t properly be written S. S. psittacina x 5. purpurea S. x courtii
flava x S. purpurea, the x reflecting the h y b r i d status S. purpurea x S. rubra S. x chelsoni
and pronounced "by" (colloquially "times") when S. alata x S. purpurea 5. x exornata
spoken. T h i s s y s t e m , although accurate and to the S. flava x S. leucophylla S. x mooreana
point, is c l u m s y in rapid c o n v e r s a t i o n or reading, e s - S. leucophylla x S. minor S. x excellens
pecially when o n e gets into hybrids c o m p o s e d of three S. leucophylla x S. psittacina S. x wrigleyana
or four or more a n c e s t o r s ! S o , the old m i s t a k e n species S. leucophylla x S. rubra S. x rearf/i
n a m e s have b e e n i n f o r m a l l y allowed (and some n e w S. fl/flfa x S. leucophylla 5. x areolata
ones have b e e n added in m o d e r n times, largely in 5. flava x S. minor S. x harperi

50/ Carnivorous Plants of the United States and Canada

S. minor x S. psittacina S. x formosa complex combinations confound even experts who
5. minor x S. rubra 5. x rehderi attempt to analyze parentage by simple inspection.
S. flava x S. rubra 5. x p o p e ; G e n e r a l l y , the best approach is to n o t e carefully w h a t
S. psittacina x S. rubra S. x gilpini species of potential p a r e n t s are n e a r b y . ( T h i s will o f t e n
5. alata x 5. rubra S. x ahlesii not w o r k very well along drainage d i t c h e s ! ) T h e n , r e -
m e m b e r i n g the tendency of Sarracenia hybrids to h a v e
an appearance intermediate b e t w e e n those of the t w o
DIFFICULT IDENTIFICATIONS p a r e n t s , o n e should try to pick out the species c h a r a c -
W i t h some experience and care, one will h a v e very teristics in the h y b r i d . T h i s can be quite interesting,
little trouble identifying species of Sarracenia in the and with some e x p e r i e n c e , you will gain perspective
field. T h e only three species that closely resemble one e n o u g h to analyze readily m o s t hybrids.
another in some ways are S. flava, S. oreophiia, and S.
alata. 5. oreophiia is completely restricted to its range
in northeastern A l a b a m a and is b e c o m i n g so rare that
the casual observer is unlikely to c o m e upon it except
in collections of live plants. T h e pale flower and s h a r p -
ly curved phyllodia are characteristic. S. flava and S.
alata can be found in the same b o g s in a few instances
around M o b i l e B a y , where their otherwise separate
ranges intersect. A careful comparison of the p h o t o -
graphs will show that S. flava has a much more p r o -
nounced and reflexed c o l u m n , a larger and flatter lid,
and a more widely flaring m o u t h . Flower differences
are also present and can be used f o r identification if
one comes upon the plants in that s t a g e : the petals of
S. flava are bright yellow and s t r a p - s h a p e d , and there
is a strong feline odor, while the petals of S. alata are
a paler yellow and more rounded, and the m u s t y odor
is far less strong. W h e n 5. alata is growing in good
sunlight, the external surface of the pitcher is more
likely to be a diffuse, pale, y e l l o w - g r e e n with fine red
veins, and in m a n y examples of the plant the u n i f o r m l y
dark red color of the inner lid and c o l u m n is distinct
from the purple splotch and coarser vein p a t t e r n s of
typical examples of 5. flava.
In larger b o g s , especially on the G u l f coastal plain,
one will surely come across h y b r i d s . S o m e of the more

51 / The Eastern North American Pitcher Plants

IV. The California Pitcher Plant (Darlingtonia
California* Torr.)
BOTANICAL NAME: Darlingtonia californica Torr. DESCRIPTION.—Darlingtonia has mainly erect to
Unacceptable synonym: Chrysamphora californica. sometimes decumbent tubular pitcher leaves which
Family Sarraceniaceae. g r o w up to 90 cm, the semidecumbent leaves tending
COMMON NAMES: California pitcher plant, cobra to be smaller. T h e pitcher leaves are n a r r o w at the bot-
p l a n t , c o b r a lily. t o m and widen to 12-15 cm in a s o m e w h a t globose
RANGE: Pacific coastal bogs and mountain slopes h o o d at the t o p . At the top of the m o d e r a t e ala is the
from Oregon to northern California. Altitude varies trap opening, which faces downward and is rather
f r o m sea level to 2800 m. large, measuring up to 3 cm. An apronlike, two-lobed,
"fishtail" appendage projects d o w n w a r d from the out-
side e d g e of the pitcher o p e n i n g opposite the ala. F r o m
a side v i e w , the w h o l e effect is that of a fancied c o b r a
with expanded hood and a rather large, protruding,
forked tongue.
W h e r e a s i n t h e g e n u s Sarracenia t h e p i t c h e r o p e n -
i n g s all t e n d t o f a c e t h e c e n t e r o f t h e r o s e t t e , t h e p i t c h -
e r s o f Darlingtonia t w i s t 1 8 0 ° i n e i t h e r d i r e c t i o n a s t h e y
g r o w , so that the pitcher openings always face a w a y
from the center of the rosette.
The pitchers are mostly pale yellow-green above
a n d a d a r k e r g r e e n b e l o w . I n full s u n l i g h t , t h e r e i s f r e -
quently m u c h red to yellow coloration of the upper
portions of the pitchers. T h e hood area has n u m e r o u s
confluent light w i n d o w s , or fenestrations. (See dis-
cussion of fenestrations u n d e r Sarracenia minor, page
38.) T h e r e a r e n e c t a r g l a n d s o v e r m u c h o f the u p p e r
external surface of the leaf, and these are especially
well developed on the tonguelike appendage.
FLOWERING SEASON: April to August, depending T h e inner m a r g i n of the pitcher opening is rolled
on altitude. into a small collar, and this feature, in addition to n u -
T R A P S E A S O N : Traps tend to remain over winter if m e r o u s d o w n w a r d - p o i n t i n g hairs lining the h o o d a n d
moderately protected. depths of the pitcher, discourages the escape of prey.

5 2 / Carnivorous Plants of the United States and Canada

T h e r e is a smooth, hairless zone between the h o o d
and the b o t t o m of the pitcher, however. Darlingtonia
has no digestive glands. It is p r e s u m e d that prey is
decomposed by microorganisms and that the nutrients
are then a b s o r b e d directly by the lining cells. T h e r e
are n o intrinsic e n z y m e s i n t h e p i t c h e r fluid, b u t t h e
presence of prey, together with certain chemical or
mechanical stimulations, does increase the secretion
o f w a t e r f r o m t h e lining cells i n t o t h e p i t c h e r c a v i t y .
T h e plant is perennial, with a long, b r a n c h i n g rhi-
zome and fibrous roots. Vegetative reproduction is
more prevalent than sexual reproduction. The nu-
merous stolons (runners) which g r o w from the rhizome
are probably m o s t responsible for the m a s s i v e prolif-
eration of Darlingtonia at suitable locations.
T h e flower h a s a tall s c a p e w h i c h a s s u m e s t h e f o r m
of a shepherd's hook at the time of anthesis (opening).
T h e r e are several b r a c t s (leafletlike blades) at i n t e r v a l s
on the scape. T h e b r a c t s are colored f r o m pale y e l l o w -
green to pink to deep red. T h e a c t i n o m o r p h i c flower
has five elongate green sepals, which project hori-
z o n t a l l y o r r e f l e x a t t h e i r b a s e s , a n d five p e n d u l o u s
crimson petals, which c o m e to a point and are closely
approximated, the result appearing to be a closed, coni-
cal c o r o l l a . N e a r t h e tip o f e a c h p e t a l , t h e l a t e r a l b o r - Fig. 4-1. A clump of Darlingtonia. Note the rotation of
ders have semicircular notches, so that, w h e n the petals the pitchers as they have grown, and the large hood.
a r e s e e n t o g e t h e r , t h e r e a p p e a r t o b e five c i r c u l a r o p e n -
ings into the corolla. and deposit pollen collected from other flowers. T h e
Inside the corolla, the large o v a r y is bell-shaped, a n d pollinator then ascends the bell-shaped ovary to the
the v e r y short style and five-lobed stellate s t i g m a p r o - flower base, w h e r e it m a y collect additional pollen.
ject d o w n f r o m t h e flat, w i d e b o t t o m s u r f a c e o f t h e As it leaves the flower, the sloping configuration of
bell. T h e t w e l v e t o f i f t e e n s h o r t s t a m e n s a r e a r r a n g e d the o v a r y tends to prevent the pollinator f r o m t o u c h -
a r o u n d the n a r r o w e r b a s e of the bell. T h i s a n a t o m i c a l ing the s t i g m a again, this time with the flower's o w n
structure serves to e n c o u r a g e cross-pollination. A pol- pollen. A l s o , t h e bell s h a p e p r e v e n t s pollen f r o m falling
linator m a y enter the flower t h r o u g h the circular o p e n - directly from the stamens onto the stigma, which is
ings in the corolla or m a y separate the tips of the largely shielded under the wide b o t t o m surface of the
petals. It will likely b r u s h o v e r the s t i g m a at this t i m e bell.

53 / The California Pitcher Plant

 T h e seeds set b y a u t u m n . T h e y are pale b r o w n and n a t u r e of the pollination process and the lack of di-
e l o n g a t e , up to 2 mm in l e n g t h . T h e b u l b o u s end of gestive enzymes, which was later confirmed by the
each seed is covered with n u m e r o u s s h o r t p r o j e c t i o n s , entomologist Frank M o r t o n Jones. She even braved a
which m a y indicate dispersion b y a n i m a l s . v i o l e n t m o u n t a i n t h u n d e r s t o r m t o sit a m o n g t h e p l a n t s
and observe that the h o o d s do indeed effectively pre-
G E N E R A L . — D a r l i n g t o n i a is quite distinct f r o m the vent the entry of rainwater into the pitchers.
eastern Sarracenia, although if is a member of the Darlingtonia grows in sphagnum bogs or in poor
same f a m i l y . Large n u m b e r s o f these plants g r o w i n g peat soils and gravel near springs and cool, fast-
in g r e a t , c o o l , g r e e n , w e s t e r n b o g s are equally as running streams. T h e geologic base rock is serpentine,
attractive as eastern pitcher p l a n t s . which has very poor nutrient value. While summer
T h e species was discovered i n 1 8 4 1 b y J . D . B r a c k - d a y s m a y b e c o m e quite w a r m , the rhizomes and roots
enridge, a b o t a n i s t assigned to an e x p l o r a t o r y e x p e d i - are a l w a y s i m m e r s e d in b o g g y slurries that are c o n -
tion i n California. The expedition was constantly stantly p e r m e a t e d with cold spring w a t e r s which sel-
threatened with hostile a t t a c k , b u t B r a c k e n r i d g e p e r - dom exceed 20°C. T h e plants occur only rarely in
sisted in w a n d e r i n g f r o m the p r o t e c t i o n of the m a i n standing-water bogs and seem to do best i n — i f not
force in order to b o t a n i z e . He c a m e upon Darlingtonia r e q u i r e — c o o l , m o v i n g w a t e r a t the r o o t level. They
and had only a f e w s e c o n d s to tear off portions of a f e w m a y thus be seen growing in seepage bogs and spring-
leaves and a seed capsule or two b e f o r e he had to rush heads, along streams and ponds, and even with mini-
b a c k to the g r o u p . B a c k in the east, the r e n o w n e d mal foothold in the snags of rapids and waterfalls.
b o t a n i s t J o h n T o r r e y i m m e d i a t e l y realized that the T h e chief thing that attracts prey seems to be the
species was n e w and t h a t it was related t o , b u t differ- gland-bearing "fishtail" hood appendage, which is
ent f r o m , Sarracenia. He f o r m a l l y described and n a m e d present in modified f o r m even in the earliest seedlings.
t h e plant f r o m the s c a n t material hurriedly a s s e m b l e d . T h e appendage attracts not only flying prey to erect
S e v e r a l y e a r s later, a specific search w a s m a d e f o r pitchers but ground prey to decumbent pitchers as
the plant, and it was f o u n d in several areas. It was well, w h e n the pitchers are bent and twisted so that
m o s t i n t e n s i v e l y and lovingly studied by a M r s . A u s - t h e l o b e t o u c h e s t h e g r o u n d . O n c e , w h i l e d o i n g field
tin, w h o g r e w up in the late n i n e t e e n t h c e n t u r y in t h e studies on the species, Frank M o r t o n Jones was carry-
area of the F e a t h e r R i v e r in C a l i f o r n i a . As a child, ing an a r m l o a d of pitchers b a c k to c a m p w h e n he n o -
she had guided s o m e of the botanical explorers a f t e r ticed a b u t t e r f l y fluttering n e a r a n d finally l a n d i n g on
Brackenridge to stands of Darlingtonia located in an the lobe of one of the pitchers, even as he was walking.
a d j a c e n t valley, which she herself had explored. A l - Like Sarracenia, Darlingtonia has several insect as-
t h o u g h not f o r m a l l y t r a i n e d , M r s . A u s t i n spent several sociates, the m o s t c o m m o n of which is the larva of a
decades studying Darlingtonia and submitting de- gnat, Metriocnemus edwardsi, which lives in the pitch-
tailed n o t e s to the r e n o w n e d eastern b o t a n i s t A s a e r liquid b u t a p p a r e n t l y d o e s n o h a r m t o the plant.
G r a y . T h e s e n o t e s remain t o this day t h e m o s t c o m - Fortunately, the California pitcher plant seems
plete and e x h a u s t i v e field s t u d y records available on destined to be with us for a long while, since at the
the species. I t w a s M r s . A u s t i n w h o d e t e r m i n e d the m o m e n t i t i s n o t s e r i o u s l y e n d a n g e r e d . M a n y g o o d Dar-

54 / Carnivorous Plants of the United States and Canada

lingtonia areas are parts of state and federal r e s e r v e s ,
and the rough, poor land and s t u n t e d , d e f o r m e d trees
of mountainous areas discourage agriculture and f o r -
estry. T h e difficulty of access to m a n y of the b o g s p r e -
vents their being overrun with visitors or easily
vandalized. H o w e v e r , even some of our m o s t r e m o t e
and primitive areas are lately suffering increased use
by weekend p i o n e e r s , and there is a g r o w i n g m a r k e t
for the species in the nursery trade.

Fig. 4-2. Art intact flower.

Fig. 4-3. A flower with two petals removed. Note the

notching of the remaining petals, the large bell-shaped
ovary, and the stamens ringing the base of the ovary.

Fig. 4-4. Close-up of the upper portion of a pitcher. One

can clearly see the fishtail appendage, the pitcher opening,
and many light windows.

55 / The California Pitcher Plant

V. The Sundews (Drosera L.)
The Genus

B O T A N I C A L N A M E : Drosera L . F a m i l y D r o s e r a c e a e . the sun. T h e stalked glands are of two main types.

COMMON NAMES: Sundew, catch-fly. (Many spe- T h o s e on the periphery of the leaf h a v e longer stalks
cies d o n o t h a v e individual c o m m o n n a m e s . ) and function mainly in entrapment. Those near the
RANGE: Mountain and coastal bogs of the Pacific c e n t e r s of b r o a d e r leaves are s h o r t e r - s t a l k e d to sessile
northwest into northern California; and, generally, in (stalkless) and function in further entrapment, but
bog locations throughout the eastern third to half m a i n l y i n s e c r e t i n g d i g e s t i v e fluids. Leaves are pro-
of North America. duced c o n t i n u o u s l y all season.
FLOWERING SEASON: Spring and summer; varies P r e y , usually v e r y small insects, is p e r h a p s lured to
somewhat with species. the trap leaves by the coloration and the sweet nectar
T R A P S E A S O N : Varies with species; most form win- s e c r e t i o n s o f the g l a n d s , o r the flying insect m a y s i m -
ter hibernacula. (See discussion below.) ply be seeking a landing platform. W h e n crawling or
alighting on a leaf, the prey b e c o m e s mired d o w n in
D E S C R I P T I O N . — T h e s e are generally perennial her- the sticky secretions, and the longer-stalked peripheral
baceous plants, although they sometimes have an an- glands then slowly b e n d in to the center of the leaf,
nual cycle and reappear t h r o u g h seed germination the placing and securing the prey in the digestive area of
f o l l o w i n g s e a s o n . T h e p l a n t s all f o r m a r o s e t t e p a t t e r n the sessile glands. The mechanism for bending the
and h a v e fibrous roots and a stem of variable length gland stalks is not u n d e r s t o o d , although electrical ac-
which m a y rise a b o v e the g r o u n d . T h e leaves are erect tivity is involved. In s o m e species a v e r y slow leaf fold-
or prostrate, depending on species and situation, and ing takes place, tending to secure the trapped prey
with the exception of one species, the stem is quite even further. This leaf folding movement is most
short. T h e leaves are of two parts: a n a r r o w , linear clearly observed in D. rotundifolia and D. intermedia.
petiole (leaf stem) of variable length, and a terminal T h e flowers are on a tall scape arising from the
blade modified into a trap. stem b e t w e e n t w o leaves. T h e f o r m of the inflorescence
T h e trapping portion of the leaf varies in shape f r o m m o s t closely a p p r o x i m a t e s a r a c e m e , since each flower
circular to linear to filiform (threadlike) in different h a s a s h o r t pedicel b e t w e e n its r e c e p t a c l e a n d t h e s c a p e ,
species. It is a l w a y s flat and has numerous stalked a s d o e s a lily o f t h e v a l l e y ; b u t i t i s c o m m o n l y r e f e r r e d
glands, mostly over the upper surface, with fewer on t o a s a s p i k e . T h e r e a r e o n a s p i k e f r o m five t o t h i r t y
the lower. T h e glands are frequently bright red and flowers lined c o n s e c u t i v e l y d o w n o n e side. T h e lowest
have a dewdroplike secretion in humid situations. flower opens for one or t w o days, then closes, and the
O f t e n , the entire leaf blade seems to be glistening with o t h e r s follow suit in n e a r l y daily succession u p w a r d
d e w — h e n c e the c o m m o n n a m e and the generic n a m e , to the tip of the spike. T h e a c t i n o m o r p h i c flowers are
from droseros, which, in Greek, means glistening in rather ordinary, the parts being mainly in multiples of

56 / Carnivorous Plants of the United States and Canada

five: five sepals, five petals, five to ten stamens, and drainage ditches in the southeast. In other places,
a five-part ovary. T h e r e are three two-lobed styles. roadside banks are so red with s u n d e w s that the plants
T h e flowers average 0.3-1.0 cm across, but the flowers m i g h t be mistaken by a passing motorist for a r a w clay
of some species m a y be as large as 2.5 c m . T h e petals surface.
m a y be white to rose pink. Petal color is n o t a reliable An interesting feature of s o m e species is the a u t u m n
w a y to differentiate species in this g e n u s , w i t h the e x - production of hibernacula, or winter buds. This occurs
c e p t i o n o f D . filiformis, w h i c h t y p i c a l l y h a s r o s e - p i n k in species that g r o w exclusively in the mountains or in
petals. the north, or in species w h o s e ranges are mainly n o r t h -
During the day, w h e n the flowers are open, c r o s s - ern but m a y extend into the southeast. T h e species of
pollination is effected by wind or small insects. If Drosera found only in the s o u t h e a s t e r n coastal plain
cross-pollination does not take place during the day, do not produce hibernacula. T h e winter bud is actually
self-pollination occurs in m o s t species as the flower a tight, s o m e w h a t spherical g r o u p i n g of leaf p r i m o r -
closes at night, thus insuring the continued p r o d u c t i o n dia (budlike y o u n g leaves), often markedly hirsute,
of seed. f r o m w h i c h t h e p l a n t will g r o w a g a i n in the s p r i n g . A
T h e seeds are b l a c k a n d elliptical a n d a r e less t h a n hibernaculum is evidently better able to withstand cold
1 mm long. Their surface patterns are consistent for weather than is an open rosette, and its formation
e a c h species, a n d in f a c t , w i t h the aid of a m i c r o s c o p e , serves to protect the plant f r o m total destruction over
o n e c a n identify t h e species by its seed. winter. After the f o r m a t i o n of hibernacula occurs, the
r e m a i n i n g leaves a n d frequently the roots die b a c k ,
G E N E R A L . — W o r l d w i d e , there are nearly a hundred leaving no trace of the plant unless one has a sharp
species of this genus. Sundews have proved quite eye for finding the w i n t e r b u d a m o n g debris.
durable and seem to withstand the rigors of environ-
mental misuse better than most other carnivorous
plants. Indeed, a disturbed site where other native
v e g e t a t i o n h a s been d e s t r o y e d p r o v i d e s an ideal o p e n -
i n g f o r t h e c o l o n i z a t i o n o f s e e d l i n g s . Drosera i s a m o n g
t h e first p l a n t s t o c o m e b a c k i n c u t o v e r s i t e s a n d a f t e r
roadside ditching and burning.
W h i l e m a n y of the N o r t h A m e r i c a n species are r a t h -
er small, some reach spectacular size. D. filiformis v.
tracyi, along the Gulf coast, for instance, has leaves
a half meter long. Even the smaller species are fasci-
nating w h e n seen in masses. Early on a bright morning
t h e i r d e w y , s t i c k y t e n t a c l e s s h i n e a n d g l i t t e r w i t h all
the colors of the spectrum. M a t s of at least one species
grow floating on the edges of, and in some places
nearly across, slow, acid streams, and the water of

57 / The Sundews
 The Species

Drosera rotundifolia L.

BOTANICAL NAME: Drosera rotundifolia L.

COMMON NAME: Roundleaf" s u n d e w .
R A N G E : In the west, from Alaska south through Brit-
ish C o l u m b i a , w e s t e r n M o n t a n a , a n d n o r t h e r n C a l i -
fornia; in the east, from Labrador and N e w f o u n d l a n d
west to the Great Lakes and south into the A p p a l a -
chians. A reported location for this species in the
C a r o l i n a coastal plain is n o t confirmed at this writing.

FLOWERING SEASON: June to September. Fig. 5-1. Drosera rotundifolia, whole plant in flower.

TRAP SEASON: Spring and summer. Forms hiber- Fig. 5-2. D. r o t u n d i f o l i a , close-up of leaf blade. The blade
nacula. is wider than long.

58 / Carnivorous Plants of the United States and Canada

D E S C R I P T I O N . — T h e plant rosette averages up to 8 Drosera linearis Goldie
cm across and becomes ungainly where it grows in
deep, coarse s p h a g n u m . T h e petioles are long, up to 4 BOTANICAL NAME: Drosera linearis Goldie.

cm. T h e leaf blade is up to 1 cm across and c a n be RANGE: From Labrador west into the Great Lakes

round but is usually wider than it is long, an i m p o r t a n t area and south to Michigan.

characteristic for identification. T h e flowers are m o r e

often white than pink.

G E N E R A L . — C h a r l e s Darwin devoted nearly half of

his book Insectivorous Plants to this species. He per-
formed numerous physiological experiments and ob-
servations and recorded his results with his usual
attention to intricate detail. T h i s is certainly o n e of
the most widespread species of Drosera and occurs
in Europe as well as in A m e r i c a .
T h e plant can usually be found in s p h a g n u m bogs,
and often coarse tufts of moss have grown up so that
the trap leaves are barely visible. If one stoops d o w n
and separates gently the strands of s p h a g n u m , the rest
of the plant is often disclosed. T h e largest and best
developed plants of this species that I h a v e ever seen
are in the N e w Jersey Pine B a r r e n s and in s o m e b o g s
in O r e g o n , w h e r e the leaf blades are d i m e - s i z e d .

F L O W E R I N G S E A S O N : June to August.
TRAP SEASON: Spring and summer. Forms hiber-
nacula.

5 9 / The Sundews
D E S C R I P T I O N . — D . linearis h a s g e n e r a l l y e r e c t l e a v e s G E N E R A L . — T h i s interesting species is restricted to
with 2 cm petioles and gland-bearing blades which cool regions of the border area between the United
g r o w up to 3 cm. T h e leaves are quite e l o n g a t e — h e n c e States and C a n a d a . It quite characteristically g r o w s in
the specific name, linearis. The flowers are usually alkaline marl b o g s , rather than in the acid situations
white. that are m o r e c o m m o n for m o s t carnivorous plants.
V e r y f r e q u e n t l y , i t i s a c c o m p a n i e d b y Sarracenia pur-
purea, w h i c h c a n g r o w i n e i t h e r a c i d o r m a r l c o n d i t i o n s
in the n o r t h , and by t w o other acid-preferring species

Fig. 5-3. D. l i n e a r i s , whole plant. Fig. 5-4. D. linearis, single leaf. The blade is long with
parallel sides.

60/ Carnivorous Plants of the United States and Canada

of Drosera, D. rotundifolia and D. anglica, which grow Drosera anglica Huds.
o n the sides a n d tops of acid, m o s s y h u m m o c k s or
BOTANICAL NAME: Drosera anglica Huds. Unac-
t u f t s s c a t t e r e d like little i s l a n d s o v e r a n d a r o u n d t h e
ceptable synonym: Drosera longifolia.
edges of the wetter, m a r l y areas. In this situation, b o t h
RANGE: In the west, f r o m the Aleutians south to
D . linearis a n d t h e a c i d - l o v i n g p a i r a r e a b l e t o g r o w
Alberta, western Montana, and into northern Cali-
in respectively suitable alkaline or acid habitats in the
fornia; in the east, f r o m Labrador w e s t into the G r e a t
s a m e b o g . P l a n t s o f D . linearis w i l l e v e n g r o w i n s h a l -
Lakes area.
low water, if it does not cover the leaves and is not
acid.
T h e matter of the plant's unique occurrence in marl
conditions has often been looked into superficially, but
the basic question of w h e t h e r the species' adaptability
t o a l k a l i n e soils i s o b l i g a t o r y o r a s e c o n d a r y a d v a n t a g e
has not been resolved. Several other people and I have
found that in culture, the seeds germinate and the
plants grow as well in acid, sand-peat soils and in
s p h a g n u m as they do in m o r e alkaline conditions, y e t
in nature, the species is only rarely found ascending
the acid, m o s s y h u m m o c k s w h i c h are frequently found
in marl bogs and which support several species of acid-
l o v i n g p l a n t s . P e r h a p s D . linearis i s u n a b l e t o c o m p e t e
with the inhabitants of the m o r e acid microhabitats
a n d t a k e s a d v a n t a g e o f its a d a p t a b i l i t y t o m a r l c o n d i -
tions where few other herbaceous plants grow and
compete for space.
T h e species is on a m a r k e d decline, particularly in
recently well-documented locations in southeastern
Michigan. The decrease seems to be correlated with
the deterioration of marl b o g s to acid conditions and
with a string of persistent rainy seasons with flooding
t o t h e e x t e n t t h a t t h e p l a n t s o f D . linearis w e r e t o t a l l y FLOWERING S E A S O N : June to August.
submerged. W h i l e the species does g r o w in shallow TRAP SEASON: Spring and summer. Forms hiber-
m a r l w a t e r s to a depth of a c e n t i m e t e r or s o , it will n o t nacula.
tolerate prolonged flooding. Other possible factors
leading to deterioration are toxic pollutants entering D E S C R I P T I O N . — T h e leaves are semidecumbent (par-
natural water systems through runoff or as lands are tially reclining) with rather long petioles reaching 3
developed; and, of course, the total drainage of a bog. to 4 cm. T h e blades are pale green with bright red,

61 / The Sundews
 stalked glands, and are longer than wide, measuring
t o 2 . 5 b y 1.0 c m . T h e f l o w e r s a r e m o s t o f t e n w h i t e .

G E N E R A L . — T h i s very attractive s u n d e w is especially

interesting to botanists because there is pretty good
evidence that it has evolved from two other contempo-
rary species, D. linearis and D. rotundifolia. The sim-
ple h y b r i d b e t w e e n t h e s e t w o , f r e q u e n t l y f o u n d w h e r e
t h e y are s y m p a t r i c , is sterile. H o w e v e r , if the n u m b e r
o f c h r o m o s o m e s o f t h e cells o f the h y b r i d e m b r y o i s
d o u b l e d t h r o u g h a n a c c i d e n t o f cell d i v i s i o n ( a m p h i -
p l o i d y ) , t h e n the flowers of the plant g r o w i n g f r o m this
e m b r y o will b e fertile, a n d t h e p l a n t will r e p r o d u c e
sexually true to species. The sterile h y b r i d and the
fertile h y b r i d ( D . anglica s p e c i e s ) look v e r y m u c h the
same outwardly, although microscopic examination
discloses l a r g e r cells in t h e a m p h i p l o i d s p e c i m e n s . In
view of the origin of the plant, some botanists would
p r e f e r t o w r i t e i t a s a h y b r i d all t h e t i m e ( D . x anglica),
rather than as a species, and they would simply note
w h e t h e r the p a r t i c u l a r plant is sterile or fertile. T h e r e
remains one v e r y perplexing p r o b l e m : given the g e o -
logic sequence of events in N o r t h A m e r i c a , h o w does
D. anglica h a p p e n t o o c c u r i n t h e w e s t a n d e v e n i n
Europe, where no D. linearis h a s e v e r b e e n r e c o r d e d ?

T h i s bright s u n d e w o c c u r s in acid s p h a g n u m b o g s ,
or on acid m o s s y h u m m o c k s in marl bogs, frequently
i n t h e c o m p a n y o f its p r o b a b l e a n c e s t o r s .

Fig. 5-5. D. a n g l i c a , whole plant.

Fig. 5-6. D. anglica. The leaf blade is somewhat oval, is

pale green, and has red glands.

62 / Carnivorous Plants of the United States and Canada

 Drosera intermedia Hayne
BOTANICAL NAME: Drosera intermedia Hayne.
R A N G E : In suitable locations over m o s t of the eastern
third of N o r t h America.

Fig. 5-7. D. intermedia, plant in flower. While the leaves

resemble those of D. anglica, the blades are somewhat nar-
F L O W E R I N G S E A S O N : June to August. rower and smaller, and there is more red pigment in the
TRAP SEASON: Spring and summer. Forms hiber- leaf tissue. One can see the longer stem formation, even in
nacula. this early summer plant.

63 / The Sundews
 Drosera filiformis Raf.
BOTANICAL NAMES: Drosera filiformis Raf. Two
f o r m s or varieties are also generally recognized within
t h e s p e c i e s : D . filiformis v . filiformis Raf. (also k n o w n
as D. filiformis v . typica W y n n e ) and D. filiformis v.
tracyi (Macfar.) Diels.
C O M M O N N A M E S : Threadleaf sundew, dew-thread.
R A N G E : As a species, f r o m C a p e C o d along the coastal
plain into southern Mississippi.

Fig. 5-8. D. i n t e r m e d i a leaf from shade-grown plant (so red

pigment of sun-grown plants would not interfere with
photographic contrast). Note the longer-stalked peripheral,
or trapping, glands and the almost sessile central digestive
glands. Several stalked glands are bent over the remnants
of small prey.

D E S C R I P T I O N . — T h i s species is unusual in that the

plant stems are quite long. In fact, the plant can reach
a height of up to 20 cm as the season progresses. T h e
left h a s m u c h the same f o r m as in D. anglica, e x c e p t
that the trapping blade is shorter and n a r r o w e r (0.5
b y 1.0 cm) and has a diffuse, d a r k red color w h e n
g r o w i n g in the open. T h e flowers are usually white.

GENERAL.—This species has the largest range of

a n y s p e c i e s o f Drosera i n the eastern part of N o r t h
A m e r i c a . In the southeastern coastal plain, the species
r e a c h e s its g r e a t e s t s i z e , a n d o f t e n t h e w h o l e p l a n t i s
deep red. It is regularly seen in m o r e wet areas, par-
ticularly on the margins of streams, ponds, and drain-
age ditches, where i t will grow into the water and
sometimes over the surface in dense mats. T h e phe-
nomenon of vegetative apomixis can be observed fre- FLOWERING S E A S O N : June.
q u e n t l y in D. intermedia. ( S e e C h a p t e r 2, p. 2 1 , f o r a TRAP SEASON: Spring and summer. Forms hiber-
discussion of vegetative apomixis.) nacula.

64 / Carnivorous Plants of the United States and Canada

DESCRIPTION—The leaves are erect with short
petioles and filiform g l a n d - b e a r i n g leaf b l a d e s . In D.
filiformis v. filiformis (typica), the leaf blades measure
to 25 cm and have bright red to purple g l a n d s ; in D.
filiformis v . tracyi t h e y m e a s u r e u p t o 5 0 c m a n d a r e
uniformly green. The hibernacula of v. filiformis
(typica) a r e b y f a r t h e m o r e h i r s u t e . T h e f l o w e r s , w h i c h
are always rose pink, are the largest of a n y variety of
Drosera a n d a r e b o r n e o n tall s c a p e s .
T h e r e a r e s o m e floral differences b e t w e e n t h e v a -
rieties. The flowers of v. filiformis (typica) measure
up to 1.5 cm across, and the outer margins of the
petals are smooth. On observing the flower with a h a n d
l e n s , o n e will n o t e t h a t t h e t w o a n t h e r l o b e s a r e j o i n e d
a t t h e tip a n d t h a t t h e s t a m e n f i l a m e n t i s p a l e r e d . I n
v . tracyi, t h e f l o w e r i s e v e n l a r g e r , u p t o 2 c m a c r o s s ,
and the petals frequently h a v e s o m e w h a t scalloped o u t -
side m a r g i n s . T h e larger a n t h e r lobes are s e p a r a t e , a n d
the stamen filament is green.

G E N E R A L . — T h i s is our largest sundew, and a stand

of them growing in a savannah or on the margin of
a bog is truly an impressive sight, especially on a d e w y
morning with the sun shining t h r o u g h the plants.
T h e smaller, red form occurs f r o m C a p e C o d south
into the N e w Jersey Pine B a r r e n s , w h e r e it is plentiful,
and it is found in disjunct locations in the e a s t e r n
Carolinas, Georgia, and a recently described location

Fig. 5-9. D. filiformis v. tracyi. A stand of the plants in a

closely cropped savannah, the glow caused by the early
morning sun striking their glandular leaves.

Fig. 5-10. D. filiformis v. tracyi, a single plant. The color is

diffusely pale green.

Fig. 5-11. Leaves of the two varieties of D. filiformis, v.

tracyi being pale green and v. filiformis (typica) having
red glands.

65 / The Sundews
 in northern Florida. T h e larger, green form grows in
the southern Gulf coastal area, where it is very c o m -
mon. The ranges of the two forms reportedly overlap
in S o u t h C a r o l i n a . In spite of this small area of s y m -
patry, the two forms have not been found in the same
stand, and a natural hybrid is not reported, although
hybrids h a v e been p r o d u c e d in the greenhouse.
As y o u will h a v e n o t e d , t h e r e is a m i n o r p r o b l e m

Fig. 5 - 1 2 . D. filiformis v. filiformis (typica). A stand of

the species in eastern North Carolina. This variety has a
red cast, which can be appreciated even at this distance.

Fig. 5 - 1 3 . D. filiformis v. filiformis (typica), a single plant

in the New Jersey Pine Barrens. The glandular character
of the leaves is clear, and they are generally reddish.

Fig. 5 - 1 4 . Flowers of D. filiformis v. filiformis (typica).

These are always rose pink, and the species has the largest
flowers of any Drosera in North America.

Fig. 5 - 1 5 . A hibernaculum of D. filiformis v. filiformis

(typica) just breaking in the early spring.

66 / Carnivorous Plants of the United States and Canada

of nomenclature with these varieties or forms. The Drosera capillaris Poir.
original name, D. filiformis, referred mainly to the
BOTANICAL NAME: Drosera capillaris Poir.
smaller, red plants. T h e n , the larger g r e e n plants w e r e
COMMON N A M E : Pink sundew.
considered by one botanist to be different enough to
R A N G E : T h e s o u t h e a s t e r n coastal plain f r o m Virginia
warrant designation as a separate species, and they
to eastern T e x a s , with a few disjunct locations in pied-
w e r e n a m e d D. tracyi. H o w e v e r , o v e r t h e y e a r s , a m a -
mont bogs.
jority of botanists have agreed that both plants are
subspecific forms of the s a m e species, so a varietal
designation has been used, and the larger, green, south-
ern plant has b e c o m e k n o w n as v. tracyi. A second
problem arose, however, in the reclassification of the
northern, red plant. T h e first r e c o r d o f a n y v a r i e t a l
designation is by Frances W y n n e , w h o in 1944 for-
mally described v. typica a s a n e w v a r i e t y . T h e r e is
no further reference until 1960, w h e n Carroll W o o d
c a s u a l l y m e n t i o n e d v . filiformis w h e n r e f e r r i n g t o t h e
northern, red form. By the usual rules of precedence,
Wynne's designation, which was formally described
f i r s t , s h o u l d b e official u n l e s s p r o p e r l y c h a n g e d ; b u t
somehow Wood made an undescribed nomenclatural
leap to v. filiformis, which has gained slightly more
p o p u l a r u s e . I c a n find n o r e f e r e n c e t o a f o r m a l d e s c r i p -
t i o n o f t h e v . filiformis d e s i g n a t i o n i n t h e 1 9 6 0 p a p e r
written by W o o d , w h o is usually meticulous in this FLOWERING SEASON: M a y to August.
r e s p e c t , n o r did W o o d m a k e a f o r m a l r e d e s c r i p t i o n o r T R A P S E A S O N : L e a v e s t e n d t o r e m a i n all y e a r . T h e r e
offer r e a s o n i n g at that time. He m a y be applying a are no winter hibernacula.
n e w e r t a x o n o m i c p r o c e d u r e w h e r e b y t h e first o f s e v -
eral subspecies, f o r m s , or varieties of a species bears DESCRIPTION.—D. capillaris is a prostrate rosette
the specific epithet. a v e r a g i n g 3 . 5 c m a c r o s s , b u t g r o w i n g u p t o 7.5 c m i n
As a rule, this species p r e f e r s a slightly drier h a b i t a t some e x t r e m e southern locations. T h e petioles are of
t h a n d o m o s t o t h e r s p e c i e s o f Drosera, a n d i t i s m o r e m o d e r a t e length, 1.0-2.5 c m , and the blades are s o m e -
o f t e n f o u n d in p e a t y , s a n d y soils t h a n in s p h a g n u m . w h a t elliptical, always longer than wide. T h e whole
D . filiformis i s e a s i l y f o u n d i n p r o p e r l o c a t i o n s , e x c e p t plant is frequently bright red w h e n growing in the
in areas where it m a y be partly obscured by surround- open. T h e r e is a prevalent notion that the flowers are
ing g r a s s a n d s e d g e s . I n t h e s e c a s e s , o n e will h a v e t o a l w a y s pink and that this provides a ready differential
make a close examination to distinguish the dewy from D. rotundifolia. Not so. The flowers are either
threadleaf f r o m a blade of grass. pink or white, the former s o m e w h a t more frequent.

6 7 / T h e Sundews
 G E N E R A L . — T h i s is undoubtedly the most ubiquitous
sundew throughout the southeastern coastal plain.
Ecologically speaking, I a l w a y s think of it as the mini-
mal c a r n i v o r o u s p l a n t , since if a likely location for
c a r n i v o r o u s p l a n t s c a n s u p p o r t o n l y o n e species i t will
probably be the hardy D. capillaris. A southeastern
g r a s s - s e d g e b o g w i t h o u t D. capillaris m u s t i n d e e d b e
in sorry straits.
A larger f o r m t h a t I h a v e f o u n d on the Gulf c o a s t
a m i d the r a t h e r o r d i n a r y f o r m s needs m o r e study.' S o
f a r , i t d o e s m a i n t a i n its m o r e r o b u s t g r o w t h h a b i t u s
w h e n g r o w n in the greenhouse under the same condi-
tions as the smaller form. My impression is that the
larger plant also grows and/or retains more leaves
than the smaller.

Fig. 5 - 1 6 . D. capillaris.

Fig. 5 - 1 7 . Flowers of D. capillaris. These are most frequent-

ly, but not always, pink.

68/ Carnivorous Plants of the United States and Canada

\
 Drosera brevifolia Pursh. are prostrate, and the trap blades taper back nearly to
the rosette center. T h e petioles are extremely short
BOTANICAL NAME: Drosera brevifolia Pursh. Pres-
to nonexistent—hence the specific name, brevifolia
ently unacceptable synonyms or additional related
(meaning short leaf). T h e end of the blade is s o m e w h a t
species: Drosera annua, Drosera leucantha.
flattened so that the leaf is cuneate ( w e d g e - s h a p e d ) .
COMMON NAMES: Shortleaf sundew, dwarf sun-
T h e entire plant is frequently red-purple w h e n g r o w -
dew.
ing in the open. T h e flower scape is uniquely g l a n d -
RANGE: In suitable locations throughout the south-
bearing, and the flowers are pink or white.
eastern United States, more commonly in the Gulf
coastal plain.
G E N E R A L . — A small n o m e n c l a t u r a l flurry o c c u r r e d in
the sixties w h e n a botanist attempted to discredit the
species n a m e brevifolia a n d t o d i v i d e t h e p o p u l a t i o n s
into at least two species, D. annua and D. leucantha.
But his arguments were not at all clear, let alone
persuasive or documented, and he m a d e the error of
relying heavily on flower and leaf color and a few
other questionable and very minor differences that
could have represented e n v i r o n m e n t a l influence. His
thesis w a s followed by a far m o r e reasoned and well-
researched paper by Carroll W o o d urging the preser-
vation of brevifolia as a species name. Wood's study

Fig. 5-18. Flower of D. brevifolia. Note the glandular scape,

which readily distinguishes this species from several others.

Fig. 5-19. D. brevifolia, the smallest species of Drosera,

has a very short petiole and a wedge-shaped leaf with a
flattened end.
F L O W E R I N G S E A S O N : April t o M a y .
T R A P S E A S O N : In good locations, the leaves remain
over winter. However, in locations that dry in the
s u m m e r , the adult p l a n t s m a y die off, a n d s e e d s will
germinate the following year. There are no winter
hibernacula.

DESCRIPTION.—This is a tiny rosette measuring

barely 2 cm across at the most. Larger plants occur
toward the west, in Louisiana and T e x a s . T h e leaves

69 / The Sundews
of the problem took four years to complete, and, dur- D. anglica and D. intermedia do have some range
ing that time, the suggested substitute n a m e s b e g a n to o v e r l a p , but this is confined to a n a r r o w a r e a in M i c h i -
appear in the literature. gan. T h e former is m o r e northern and has a very short
This is o u r tiniest s u n d e w , a n d one m u s t look close- s t e m a n d wider a n d longer leaf blades, w h i c h are g r e e n
l y e v e n t o find i n d i v i d u a l s a n d t h e n d i f f e r e n t i a t e t h e m with bright red glands. D. intermedia i s m o r e s o u t h e r n
from D. capillaris, with which they often grow. D. a n d has a m u c h longer s t e m in older plants, a smaller
brevifolia is often present in glittering masses on a leaf, a n d red p i g m e n t in the plant tissue w h e n g r o w -
r o a d s i d e b a n k o r s a n d y flat. C l o s e r i n s p e c t i o n o f s u c h ing in the open.
areas discloses the individual plants. D. brevifolia B y t h e w a y , a r a r e n a t u r a l h y b r i d o f D . filiformis v .
seems to prefer a drier habitat than m o s t of our other filiformis (typica) and D. intermedia, which had been
species of Drosera. described in the N e w Jersey Pine Barrens, w a s recently
rediscovered. Its a p p e a r a n c e is intermediate b e t w e e n
those of the t w o parents, and plants studied thus far in
DIFFICULT IDENTIFICATIONS the g r e e n h o u s e a p p e a r to be sterile. T h e hybrid is s o m e -
I think the b i g g e s t difficulty m o s t people h a v e is the times known as D. x hybrida. D. rotundifolia x D.
differentiation of D. capillaris from D. rotundifolia. anglica (D. x obovata) is not infrequently found in
First o f all, f o r g e t t h e c o m m o n e r r o r o f f l o w e r c o l o r . northern bogs where the t w o species are sympatric.
Second, D. rotundifolia i s f o u n d i n m o r e n o r t h e r n a n d T h e leaf f o r m is intermediate between those of the
mountainous regions, whereas D. capillaris i s l o c a t e d t w o p a r e n t s a n d m a y b e difficult t o d i s t i n g u i s h f r o m
in the southeastern coastal plain (except for a few t h a t o f D . anglica u n l e s s t h e t w o a r e c o m p a r e d s i d e b y
bogs in piedmont Carolina, where I have found the t w o side. F i n a l l y , D. rotundifolia x D. intermedia h a s been
together). Third, and most important, the trap portion r e p o r t e d in N e w Jersey, the usual rule of intermediacy
of the leaf of D. rotundifolia i s most often wider than causing the leaf blades to a p p e a r almost round.
long, but occasionally perfectly round in younger
plants or in n e w spring leaves, while the blade of D.
capillaris is longer than wide.
D. brevifolia can be confused with the above two
species, especially when they a r e all seedlings. The
d w a r f s u n d e w has a very short petiole and a w e d g e -
shaped leaf blade tapering nearly b a c k to the rosette
center. T h e easiest identification m a r k e r for the begin-
ner is the scape of D. brevifolia, which is gland-
bearing, whereas the other two species with which it
m a y be confused have smooth scapes. Also the purple-
red color of the plants of D. brevifolia i s d i s t i n c t i v e
f r o m t h e l i g h t e r r e d o f D . capillaris, b u t t h e d i f f e r e n c e
is subtle and requires experience to discern.

70 / Carnivorous Plants of the United States and Canada

VI. The Butterworts (Pinguicula L.)
The Genus

BOTANICAL NAME: Pinguicula L. Family Lenti- together at their bases). T h e three lower petals f o r m a
bulariaceae. l o w e r lip w i t h o r w i t h o u t p a r t i a l d i v i s i o n i n t o l o b e s ,
COMMON NAME: Butterwort. a n d t h e t w o u p p e r p e t a l s f o r m t h e u p p e r lip. N e a r its
R A N G E : M e m b e r s o f t h e g e n u s o c c u r all a c r o s s t h e base, the corolla n a r r o w s into a tube of cylindrical
northern half of N o r t h America and extend d o w n into s h a p e , a n d this t e r m i n a t e s in a spur, w h i c h is e v e n n a r -
the coastal plain in the s o u t h e a s t a n d into northern rower and of variable length. Originating from the
California in the west. i n n e r s u r f a c e o f t h e l o w e r lip a n d a t l e a s t p a r t i a l l y
F L O W E R I N G S E A S O N : Varies with the species; gen- covering the e n t r a n c e to the corolla tube is a slightly
erally, early spring into early s u m m e r . bulging structure, called a palate, which is bearded
T R A P S E A S O N : T h e plants native to southern areas
retain their leaves o v e r w i n t e r ; those native to n o r t h - Fig. 6-1. Pinguicula vulgaris. These plants are showing the
ern and western areas f o r m hibernacula (winter b u d s ) . formation of early autumn hibernacula in their centers. The
general plant structure is evident.

D E S C R I P T I O N . — T h e plant is a rosette with stalkless Fig. 6-2. The flower of P. vulgaris.

leaves, the older ones lying prostrate, and the y o u n g e r
ones nearly so. T h e thin leaves are elongate and n a r -
r o w m o r e or less to a blunt point at the free end. T h e y
a r e g e n e r a l l y flat o n t h e m a i n s u r f a c e , b u t t h e m a r g i n s
are variably rolled in different species. T h e s u r f a c e of
the leaf is studded with nearly microscopic sessile
(stalkless) g l a n d s , w h i c h i m p a r t a fine, p e b b l e d t e x -
ture. T h e leaf feels g r e a s y to the t o u c h — h e n c e the
n a m e , f r o m t h e L a t i n w o r d pinguis, m e a n i n g fat, and
t h e s u f f i x -ula, m e a n i n g little o n e . T h e plant is pale
yellow-green in most species, but reddish in one. T h e
roots are fibrous and brittle, extending 2 - 6 cm into
the ground.
T h e flower scape is often gland-bearing, is 4 - 2 0 cm
long, and supports the single flower at the top. M u l t i -
ple s c a p e s a p p e a r s u c c e s s i v e l y d u r i n g the e a r l y g r o w i n g
season. T h e flower is z y g o m o r p h i c , and the corolla is
at least partly s y m p e t a l o u s (i.e., the petals are fused
(hairy). This m a y be exserted (projecting outward be- Typical Flower of Lentibulariaceae
y o n d the flared corolla surface) or not, depending on Reproductive Structures
the species. T h e hairs on and a r o u n d the palate and in
the tube of each species are seen to h a v e a distinctive
structure when they are examined under the micro-
scope.
T h e s t a m e n s a n d pistil h a v e a u n i q u e a r r a n g e m e n t
characteristic of the family. T h e y are located deep in
the tube. T h e stamen filaments are thick, stocky, and
curved, and each is capped by a spherical anther with
yellowish pollen. T h e r e are t w o s t a m e n s placed n e x t
to each other in the same plane, so that the curvature
of the filaments nearly causes the anthers to touch.
Behind the bases of the s t a m e n s and attached to the
receptacle is the spherical ovary with a very short
style. T h e stigma is bilobed and modified. T h e pos-
terior lobe is g r e a t l y r e d u c e d ; the anterior lobe is flat-
tened and somewhat elongated, so that it hangs over
t h e a n t h e r s like a veil or a p r o n . T h e u p p e r s u r f a c e of
the stigma is covered with a sticky material to hold
pollen a n d s u p p o r t its g e r m i n a t i o n .

The flower tends to encourage cross-pollination

r a t h e r t h a n selfing. T h e p o l l i n a t o r e x t e n d s its f o r e b o d y
deep into the corolla tube and, as it does so, deposits
a n y pollen f r o m another flower on the anterior stigma
l o b e . A s t h e p o l l i n a t o r w i t h d r a w s , i t t e n d s t o lift t h e
apronlike anterior stigma lobe upward (because of
friction and a tight fit), a n d e x p o s e the t w o a n t h e r s , so
that it m a y pick up pollen f r o m that flower without
depositing it on the upper surface of the anterior stig-
ma lobe, w h i c h has been turned back against the wall
of the corolla tube. (See drawing at right.)
T h e trapping m e c h a n i s m is quite simple. Small prey
landing or crawling on the u p p e r surface of the leaf
become mired down in the glandular secretions and
are held fast until digestion a n d a b s o r p t i o n take place.
During the active trapping season, the rolled edges of

72 / Carnivorous Plants of the United States and Canada

 to several other herbs growing in the same habitat. In
m a n y i n s t a n c e s , t h e n e w n a t u r a l i s t will h a v e difficulty
s p o t t i n g Pinguicula r i g h t a w a y . In late spring, h o w e v e r ,
t h e c o l o r f u l little f l o w e r s , a t o p t h e i r tall, f r a g i l e s c a p e s ,
nod in the breeze, and the plants are then m u c h m o r e
obvious, especially when massed. The color of the
flowers and the patterns of venation, if any, are c h a r a c -
teristic f o r m o s t s p e c i e s , a n d w e will s t r e s s t h e m . U n t i l
one gains experience, the vegetative part of the plant
looks very m u c h the s a m e f r o m species to species.
N o t only are the plants small, vegetatively incon-
spicuous, and perhaps not as interesting to most people
as Sarracenia, Dionaea, and Drosera, but many people
a r e repelled b y the cool, g r e a s y feel o f the leaves. T h i s
Fig. 6-3. The flower of P. planifolia, opened by removing
texture is due to the rather fragile, almost weakly suc-
the lower lip. One can clearly see the apronlike lobe of the
anterior stigma overhanging the partially exposed anthers culent nature of the leaf tissue, in addition to the glan-
atop the two curved, stout stamen filaments in the tube. dular secretions on the upper surface.
Near the right side of the picture is a portion of the spur. I k n o w of one m i n o r e c o n o m i c use f o r Pinguicula.
People in the countries of northern Europe have mixed
the leaf tend to curl in s o m e species, b u t s e l d o m to the leaves or leaf e x t r a c t s of certain species with milk
the e x t e n t o r w i t h sufficient s p e e d t h a t t h e y a c t u a l l y in order to curdle it a n d p r e p a r e a pudding dish m u c h
engulf the prey, as has been suggested in the past. like j u n k e t o r y o g u r t . B e y o n d this, t h e p l a n t s will h a v e
Studies by scanning electron microscopy have re- to be accepted a n d studied on their o w n merits. A n d
c e n t l y s h o w n t h a t t h e Pinguicula l e a f h a s t w o k i n d s o f they are fascinating plants.
glands on the upper surface, these being randomly Natural or artificial hybrids of the southeastern
intermingled. There are stalked glands that perhaps species have not been found. My own preliminary
are m o r e i m p o r t a n t in c a p t u r e , a n d sessile g l a n d s t h a t experiments with m a n y crosses s h o w that seeds are
possibly are more active in digestion. Absorption produced but do not germinate. T h e r e are reports of
probably takes place at the bases of the digestive hybrids between the species of Pinguicula that form
glands. hibernacula, but these are mainly found in other
countries.
GENERAL.—You will note the poverty of common
n a m e s f o r t h e s e little p l a n t s a s y o u r e a d a l o n g . T h i s
clearly indicates the small regard and interest that h a s
been accorded Pinguicula, except in a few scholarly
instances. T h e plants are rather inconspicuous outside
the flowering season, and there is a loose r e s e m b l a n c e

7 3 / The Butterworts
 The Species FLOWERING S E A S O N : June to August.
T R A P S E A S O N : Forms winter hibernacula, frequent-

Pinguicula vulgaris L. ly with smaller off-budding basal hibernacula, or g e m -

m a e , w h i c h will g r o w i n t o y o u n g p l a n t s i n the spring.
BOTANICAL NAME: Pinguicula vulgaris L. There
is a likelihood that a p r o p o s e d split of w h a t is c o m - D E S C R I P T I O N . — I n the northeastern range, the r o -
monly known as P. vulgaris in the west will even- settes m e a s u r e up to 11 c m . T h e leaves are pale yellow-
tually be accepted, resulting in another species, P. green and are rather wide, with somewhat irregular
macroceras Link, with three varieties: v. macroceras, m a r g i n s and minimal m a r g i n a l curling. T h e flower is
v. microceras (Cham.) Casper, and v. nortensis (un- violet with pale to white patches t o w a r d the tube. T h e
published at press time). See discussion below and c o r o l l a a v e r a g e s 1.1 c m a c r o s s , a n d t h e u p p e r lip i s
m a p of outlines of ranges. smaller than the lower. The beard is rather weak,
COMMON N A M E : Butterwort. being a confluent grouping of whitish hairs in the
R A N G E : Northern boreal (subarctic) region south to throat of the tube, and it is not exserted. T h e spur
the Great Lakes and northern California. averages 5-7 mm and is sometimes notched or double.
T h e consistent variation of plants in the west has
p r o m p t e d the concept of a separate species, which is
strengthened by the fact that, in the areas of overlap
in the far n o r t h w e s t , the plants tend to maintain their
separate characteristics. Generally, the rosettes of the
variant plants are smaller (5-9 c m ) , and the flowers
l a r g e r (1.5—1.8 c m ) . T h e m a i n d i a g n o s t i c d i f f e r e n c e i s
in the spur, w h i c h is longer in the west (6-11 c m ) , and
a l s o i n t h e l o w e r lip l o b e s , w h i c h a r e l a r g e r a n d t e n d
to overlap or at least touch. Flower coloring is about
the same. This plant has been proposed as a separate
species, P. macroceras, b y some botanists. The picture
is complicated by still further variation within the
putative n e w species. In the Aleutians, there is a variant
with a very short spur (1-3 c m ) , but with the larger
corolla. This plant is further designated v. microceras,
a n d t h e l o n g e r - s p u r r e d , m o r e s o u t h e r n p l a n t , v . mac-
roceras. T h e n , i n t h e f a r s o u t h e r n e n d o f t h e w e s t e r n
range, in Del N o r t e County, California, populations
were found with the long spur, but with smaller and
well-separated corolla lobes more like those of the
e a s t e r n P . vulgaris. T h e d e s i g n a t i o n o f t h e s u b s p e c i e s

74 / Carnivorous Plants of the United States and Canada

 m u c h the importance of such " b e h a v o r i a l " studies in
investigating the variations of plants. They are too
often overlooked by even the m o s t experienced bot-
anists, but they must be done no matter what the
cost in time and frustration. Third, there are as yet
not even elementary studies in genetic crossing. Final-
l y , t h e c h r o m o s o m e n u m b e r o f t y p i c a l e a s t e r n P . vul-
garis is known to be 2n=64, but no chromosome
counts have been done on a good cross section of
plants throughout the western range, except for one
plant of the proposed nortensis group, and this is
2n=32.
Fig. 6-4. Flower of plant from the " m a c r o c e r a s " range, Del E v e n t u a l l y , a f t e r all t h e s e r e q u i r e d s t u d i e s a r e c o m -
Norte County, California. p l e t e , t h e s e p a r a t i o n o f t h e l o n g - a c c e p t e d P . vulgaris
taxon m a y be acceptable without a doubt, and if so,
or variety nortensis (after the county) may be pro- the r e a d e r s of this b o o k will be p r e p a r e d . B u t at the
posed for these populations, which have also now moment, there is far too m u c h w o r k to be done to be
been found in immediately neighboring O r e g o n . dogmatic on the issue.
Japanese botanists, as well as many botanists in
t h i s c o u n t r y , h a v e b e e n m a i n l y c o n t e n t t o c o n s i d e r all G E N E R A L . — I n the east, this species g r o w s along the
t h e s e n o r t h e r n P a c i f i c p l a n t s a s v a r i e t i e s o f P . vulgaris. rocky and gravelly shores of lakes and streams in a
H o w e v e r , there is a strong m o v e m e n t afoot to separate thin layer of peat, or in and along the edges of both
the plants as mentioned above. U n f o r t u n a t e l y , data is s p h a g n u m and m a r l b o g s , apparently able to get along
far f r o m complete, and claims for such separation are, well in several kinds of habitats. In the winter, o n e
at publication time, premature. M o s t of the original can often see h u n d r e d s of hibernacula and g e m m a e
studies were done on herbarium material, which we floating loose a m o n g the flotsam of lakeshore slack-
are finding wholly inadequate for investigating the w a t e r in quieter inlets. This is an apparently helpful
f r a g i l e Pinguicula. I n f a c t , a m o n o g r a p h b y a E u r o p e a n m e c h a n i s m f o r d i s p e r s a l . I n t h e w e s t , t h e "macroceras"
botanist, w h o has studied the herbarium sheets and plants tend to prefer a narrower habitat; they g r o w
is p r o m o t i n g the n e w species designations, features a in seeps on m o s s y outcrops of serpentine rock, often
large number of photos of pressed plants and not more partially shaded, along the m a r g i n s of small springs
t h a n a h a l f - d o z e n live o n e s ! N o t to h a v e s t u d i e d the and bogs which often dry s o m e w h a t in the summer.
live plants more thoroughly is, in this day, totally In the far northwest, the plants are often seen on m o s -
inadequate. sy sphagnum h u m m o c k s and in open gravelly seeps.
Second, the all-important studies of reciprocal t r a n s - N o r t h e r n California plants with pale red leaves h a v e
p l a n t s as well as g r e e n h o u s e s t u d i e s in h o m o g e n e o u s occasionally been found recently growing interspersed
environments have not been done. I cannot stress too among typical green-leafed forms.

75 / The Butterworts
 Pinguicula pumila Michx.
BOTANICAL NAME: Pinguicula pumila Michx.
RANGE: T h e southeastern coastal plain f r o m N o r t h
Carolina into east T e x a s , but rare except in the Gulf
area.

Fig. 6 - 5 . The flower of P. pumila. Note that the corolla

does not appear fully expanded (a common occurrence in
this small species) and that the yellow heard is not ex-
serted.

FLOWERING SEASON: April to May.

T R A P S E A S O N : N o hibernacula are formed.

D E S C R I P T I O N . — M o s t often, the rosettes are barely

1.5 c m a c r o s s , b u t o c c a s i o n a l l y t h e y r e a c h 2 c m . T h e
leaves are pale green a n d h a v e pointed tips. T h e leaf
edges are sharply rolled. The 1.0-1.5 cm flower is
most often white but ranges to purple, yellow, or pink
in Florida. T h e pale yellow beard is not exserted.

G E N E R A L . — This is our tiniest Pinguicula, barely

discernible even w h e n in flower. It tends to g r o w in
moist, sandy places that dry s o m e w h a t — b u t not c o m -
pletely—during the summer.

76 / Carnivorous Plants of the United States and Canada

 Pinguicula lutea Walt.
BOTANICAL NAME: Pinguicula lutea W a l t .
C O M M O N N A M E : Y e l l o w b u t t e r w o r t (referring t o
the color of the flower).
R A N G E : T h e southeastern coastal plain f r o m N o r t h
Carolina to Louisiana.

F L O W E R I N G S E A S O N : February to M a y .
T R A P S E A S O N : N o winter h i b e r n a c u l a .

D E S C R I P T I O N . — T h e s e are pale green rosettes r a n g -

ing 5 - 1 5 cm a c r o s s , the larger ones tending to occur
further south. T h e leaf edges are sharply rolled, and
the leaf tips are pointed. T h e large ( 2 . 5 - 3 . 5 cm) flower
is a brilliant yellow. T h i s is the only large species in Fig. 6-6. P. lutea. The bright yellow flower is distinctive.

our area with a regularly yellow flower.

GENERAL.—A stand of P. lutea in flower in the

earliest spring is a pretty sight and will n o t easily be
missed. T h e s e plants grow in open, d a m p , sandy places
that are shaded over to some degree in later s u m m e r
by taller herbs and g r a s s e s .

77/The Butterworts
 Pinguicula caerulea Walt. Pinguicula planifolia Chapm.
BOTANICAL NAME: Pinguicula caerulea Walt. BOTANICAL NAME: Pinguicula planifolia Chapm.
COMMON NAME: Violet b u t t e r w o r t (referring t o R A N G E : T h e gulf coastal plain f r o m the mid-Florida
the color of the flower). p a n h a n d l e w e s t to Louisiana.
R A N G E : T h e s o u t h e a s t e r n coastal plain f r o m N o r t h F L O W E R I N G S E A S O N : M a r c h t o April.
Carolina to the middle p a n h a n d l e of Florida. T R A P S E A S O N : N o w i n t e r hibernacula.

F L O W E R I N G S E A S O N : February to M a y .
T R A P S E A S O N : N o winter h i b e r n a c u l a .

DESCRIPTION.—The rosettes are pale green and

measure 5 - 1 0 c m across. T h e leaves have sharply
rolled edges and are pointed. T h e scapes are tall (to
2 0 c m ) , and the large 2 . 5 - 3 . 0 c m corollas are violet
with p r o m i n e n t v e n a t i o n of a deeper violet. T h e palate
beard is exserted and colored g r e e n i s h yellow to c r e a m .
Fig. 6-7. The flower of P. caerulea. Venation is very prom-
G E N E R A L . — I n the vegetative stages o f later s u m m e r , inent in this large flower. You can just see the top of the
it is practically impossible to tell this species f r o m P. anterior stigma lobe down in the tube entrance.
lutea, with which it sometimes g r o w s . But in the spring,
Fig. 6-8. The flower of P. planifolia, with prominent ex-
the deep-veined, violet flowers of P. caerulea certainly serted beard, pale purple color, and deeply incised corolla
contrast with the yellow flowers of the o t h e r species. lobes.

78 / Carnivorous Plants of the United States and Canada

D E S C R I P T I O N . — T h e large rosettes measure to 15 cm Pinguicula primuliflora W o o d & Godfrey
across, and the older leaves are flat and have o n l y
BOTANICAL NAME: Pinguicula primuliflora Wood
slightly rolled edges. T h e leaves are mostly a dull red
& Godfrey.
to purple, although green races are c o m m o n . T h e tall
R A N G E : G u l f coastal plain f r o m the w e s t e r n Florida
scape bears a 3 c m , violet, unveined flower, which
p a n h a n d l e into s o u t h e r n M i s s i s s i p p i .
tends to be darker colored around the tube e n t r a n c e . A
prominent characteristic is that the lobes of the corolla
are deeply incised to at least half their l e n g t h , so t h a t ,
at a glance, it appears as if the corolla has ten lobes
instead of five. T h e palate beard is exserted and bright
yellow.

G E N E R A L . — In contrast to the previously described

species of the southeastern coastal plain, this and the
next two species g r o w in c o n s t a n t l y wet a r e a s , s o m e -
times completely s u b m e r g e d in water for periods of
time. T h e distinctly purple, unveined flower of P. plani-
folia, with its deeply incised corolla l o b e s , m a k e s f o r
easy identification in the spring. W h e n the plant is not
in flower, r e m e m b e r that it is the only species with red-
dish leaves. But different races and s h a d e - g r o w n plants
of P. planifolia have green leaves, and in such cases
there will be some difficulty in distinguishing n o n - F L O W E R I N G S E A S O N : February t o April.
flowering plants f r o m the next two species. T R A P S E A S O N : N o winter hibernacula.

Fig. 6-9. P. planifolia. The leaves have a reddish color. D E S C R I P T I O N . — T h e green rosettes g r o w u p t o 1 5
cm a c r o s s , and the leaf edges are m o d e r a t e l y rolled.
T h e tall scape supports a 2 . 5 - 3 . 0 c m , very pale blue
to violet flower with a white ring a r o u n d the t u b e
e n t r a n c e . T h e external surface of the tube and spur is
bright yellow. T h e palate beard is yellow and e x s e r t e d .

G E N E R A L . — T h i s species also p r e f e r s wet areas. It is

o f t e n f o u n d in and on the edges of slowly m o v i n g
s t r e a m s , w h e r e it g r o w s on h u m m o c k s of s p h a g n u m ,
m o s t l y in the shade. T h e flower is u n i q u e , as are the
flowers of all the southern species.
 T h e r e is a n o t h e r peculiarity of this plant that is
shared to a far lesser degree by the o t h e r two princi-
pally G u l f coastal species, P. planifolia and P. ionantha.
S m a l l vegetative buds frequently sprout f r o m the
tips of older leaves in late s u m m e r . T h e s e can be seen
growing f r o m decaying leaves or ringed w i t h o t h e r
plantlets of v a r y i n g ages a r o u n d a larger " m o t h e r "
plant.

Fig. 6 - 1 1 . P. primuliflora. There is a new plant bud arising

from the end of an older leaf.

Pinguicula ionantha Godfrey

BOTANICAL NAME: Pinguicula ionantha Godfrey.
R A N G E : A very n a r r o w range in the mid-panhandle of
w e s t e r n Florida.
F L O W E R I N G S E A S O N : February t o April.
T R A P S E A S O N : N o winter hibernacula.

Fig. 6 - 1 0 . The flower of P. primuliflora. The corolla is rose

pink to violet with a white center.

80 / Carnivorous Plants of the United States and Canada

D E S C R I P T I O N . — T h e flat, pale green rosettes m e a - Pinguicula villosa L.
sure up to 15 cm across. T h e r e is minimal rolling of
BOTANICAL NAME: Pinguicula villosa L.
the somewhat irregular leaf m a r g i n s . T h e tall scape
R A N G E : N o r t h A m e r i c a n boreal (subarctic and arctic)
supports a 2 cm white to e x t r e m e l y pale violet flower
region f r o m A l a s k a across e x t r e m e n o r t h e r n C a n a d a .
with a ring of deeper violet around the tube e n t r a n c e .
T h e spur and external tube are olive to dull yellow.
T h e palate beard is bright yellow and exserted.

G E N E R A L . — T h i s species g r o w s in very wet p e a t y or

sandy places and in shallow w a t e r , o f t e n a c c o m p a n i e d
by P. planifolia. It has an e x t r e m e l y small range and
is therefore s o m e w h a t endangered.

F L O W E R I N G S E A S O N : June to August.
Fig. 6-12. The flower of P. ionantha. This flower is white
T R A P S E A S O N : Forms w i n t e r h i b e r n a c u l a .
to very pale violet, with a darker ring in the center.
D E S C R I P T I O N . — P . villosa is a small r o s e t t e , b a r e l y
2 - 3 c m a c r o s s , with pale g r e e n , s o m e w h a t o b o v a t e
leaves. T h e scape characteristically has n u m e r o u s long
plant hairs over the lower p o r t i o n ; h e n c e the specific
epithet. T h e flower, w h i c h is about 1 cm a c r o s s , is
colored pale blue to violet (with a white f o r m recorded)
and has fine yellow v e n a t i o n and a w e a k b e a r d . T h e
spur averages 0.5 cm and is s o m e w h a t conical.

81 / The Butterworts
G E N E R A L . — T h i s plant g r o w s on sphagnum
and h u m m o c k s in the e x t r e m e n o r t h e r n part of the
c o n t i n e n t , w h e r e it is well adapted to the subarctic
climate. We h a v e not b e e n able to cultivate this plant
f o r protracted periods, p r o b a b l y b e c a u s e it requires
cool temperatures and some a p p r o x i m a t i o n of
long, cool s u m m e r days and the d a r k , very frigid w i n -
ters of P. villosa's natural h a b i t a t .
DIFFICULT IDENTIFICATIONS
C e r t a i n l y w h e n t h e y are in f l o w e r — a n d sometimes
w h e n they are out of f l o w e r — t h e r e will be little diffi-
culty in separating these species. R e f e r e n c e to the
p h o t o g r a p h s will s h o w floral differences at a glance.
W h e n the plants are considered regionally, identifica-
tion is o f t e n m o r e simplified.
In the area of the n o r t h e a s t e r n b o r d e r b e t w e e n the
U n i t e d S t a t e s and C a n a d a , there is only P. vulgaris.
O u r w e s t e r n readers will have to c o n t e n d w i t h the
82 / Carnivorous Plants of the United States and Canada
tufts "macroceras" h y p o t h e s i s , if it is eventually established.
Boreal b o t a n i s t s m a y run into P. vulgaris, the "macro-
ceras" s u b g r o u p s , and P. villosa. W h e n the plants are
in flower, identification will be e a s y , and the rounder
leaves and h a i r y scape of P. villosa are also c h a r a c t e r -
the istic of that species.
O n the A t l a n t i c coastal plain o f the Carolinas, there
are only P. lutea, P. caerulea, and P. pumila, all with
very distinctive flowers. Even out of flower, a s t o c k y ,
tiny rosette is m o s t likely P. pumila, which is generally
u n c o m m o n in this area. Look f o r a maturing seed cap-
sule to be sure that the plant is not a juvenile of one
of the o t h e r t w o s p e c i e s ! P. lutea and P. caerulea are
common.
In the G u l f coastal region, especially upper Florida,
there is the widest range of species, and identification
by flower will be n e c e s s a r y in m o s t i n s t a n c e s , except
w h e n the red leaves of P. planifolia and the peripheral
buddings of P. primuliflora are present.
VII. The Bladderworts (Utricularia L.)
Family Lentibulariaceae
INTRODUCTION
We will depart f r o m the f o r m a t of the preceding
chapters for several reasons. First of all, the t a x o n o m y ,
b i o l o g y , and range demarcations of the bladderworts
are not as well k n o w n and understood as those of the
other genera of carnivorous plants. T h e r e are no d e -
tailed botanical m o n o g r a p h s on the A m e r i c a n species,
although at least one is k n o w n to be f o r t h c o m i n g .
S e c o n d , the various species of Utricularia excite f a r
less popular interest than the larger terrestrial c a r n i v o -
rous species. Indeed, if it were not for their annual
flower displays, these rather m i n u t e plants could easily
be passed over entirely by the n o v i c e , even t h o u g h
they are far more ubiquitous than any of the o t h e r
carnivorous genera. T h e r e is hardly a salubriously
b o g g y place that does not support at least o n e , and
more likely several, species of Utricularia. C l e a r l y , the
genus requires and deserves far m o r e study and at-
tention.
In this chapter we will not have range m a p s , and
there will be only representative p h o t o s of a b r o a d
cross section. D e s c r i p t i o n s will be far m o r e b r i e f ,
mentioning a few salient points that should be suffi-
cient to differentiate closely related species to the e x -
tent that we understand t h e m at present. T h e s e will
serve for easiest and most rapid identification. M a n y
other points of difference will be omitted f o r the s a k e
of clarity and to eliminate excessive technicality and
controversy at this point, although we recognize their
importance to the serious botanist. For these latter
readers we suggest consulting the f e w available r e f -
erences.
83 / The Bladderworts
T h e interested reader will find that a h a n d lens of
good quality or e v e n a microscope eyepiece is required
f o r close e x a m i n a t i o n and identification o f m a n y o f
these plants.
GENERAL DESCRIPTION OF THE GENUS
T h e species of Utricularia inhabiting N o r t h A m e r i c a
are either aquatic or terrestrial, the f o r m e r f o u n d as
strands or m a t s of plants floating in quiet, acid ponds
and b o g - a s s o c i a t e d w a t e r s . T h e terrestrials grow m o s t
c o m m o n l y in d a m p , s a n d y , acid soils, with the m a i n
parts of the plants at or b e l o w ground level. W i t h o u t
m i n u t e e x a m i n a t i o n of t h e soil in the latter c a s e , o n l y
annual f l o w e r i n g signals the plant's presence. T h e t e r -
restrials also g r o w in s p h a g n u m m a t s and h u m m o c k s ,
w h i c h are s o m e t i m e s quite w e t , and t h e y can c o m -
m o n l y b e f o u n d i n m a r l b o g situations. M o s t o f the
species, aquatic and terrestrial, are frequently f o u n d
in sand and peat m u c k or s p h a g n u m slurries, an i n -
termediate h a b i t a t . T h i s capability of a biphasic h a b i -
tat is an advantage f o r survival, since ponds w i t h
primarily aquatic species o f t e n partially dry out in the
s u m m e r , and m a n y areas w h e r e terrestrials g r o w flood
during rainy s e a s o n s .
T h e f o r m of the plant is a rootless, b r a n c h i n g , green
o r b r o w n stem 0 . 1 - 3 . 0 m m i n t h i c k n e s s , f r o m w h i c h
arise whorls of finer green b r a n c h e s , w h i c h are s o m e -
times divided and almost f e a t h e r y , and w h i c h usually
bear tiny b u l b o u s traps. P r o p e r l y , the traps themselves
are the l e a v e s , b u t no one quite k n o w s w h a t to call the
trapless b r a n c h i n g s w h i c h s o m e t i m e s p r o j e c t up out
of the soil or w a t e r in leaflike f a s h i o n . T h e s e h a v e b e e n
called leaves or photosynthetic organs or branches, t e r i s t i c s o f t h e f l o w e r s o f Utricularia. R e f e r e n c e t o t h e
each n a m e adequately implying the supposed function. discussion of the flowers of Pinguicula, pp. 71-72,
T h e rootless, branching stem m a y reach a length of will be h e l p f u l .
3 m, or even m o r e in the case of some aquatics. The aerial flowers of Utricularia are borne on an
The traps or bladders have a bulbous form, and o f t e n tall, n a r r o w s c a p e , and they n u m b e r f r o m o n e
t h e y r a n g e i n s i z e f r o m 0.5 t o 3 . 0 m m a c c o r d i n g t o a g e to fifteen or e v e n m o r e , depending on the species. T h e
a n d s p e c i e s . T h e y a r e a t t a c h e d t o t h e finer b r a n c h i n g s s c a p e m a y h a v e a l o n g its c o u r s e a n d a t t h e b r a n c h i n g s
by a n a r r o w stalk at the b o t t o m of the bladder. of pedicels o n e or several m i n u t e , leaflike a p p e n d a g e s
As mentioned above, m a n y of the terrestrial species called b r a c t s , bracteoles, or associated scales. T h e n u m -
h a v e n a r r o w , flattened, pointed, green leaflike struc- ber, color, f o r m , and location of these structures can
tures that have almost the appearance of seedling grass- be important taxonomically.
blades. T h e s e arise f r o m the stem or the base of the T h e flower itself is z y g o m o r p h i c ( t w o - l i p p e d ) , as in
scape a n d project 1-5 m m a b o v e g r o u n d level. T h e y Pinguicula, b u t i t t e n d s t o b e m o r e f l a t t e n e d , a n d t h e
apparently have a photosynthetic function. s i z e a n d s h a p e o f t h e s p u r a r e m o r e v a r i a b l e . I n Utricu-
Since Utricularia a n d Pinguicula a r e m e m b e r s of the laria, the palate more completely obstructs the tube,
s a m e family, their flower structures are basically quite a n d i s h a i r l e s s . T h e r e a r e fine r e d o r b r o w n r e t i c u l a t e
similar, s o w e will p o i n t o u t o n l y s o m e specific c h a r a c - lines on the palates of several species.

Fig. 7-1. U. m a c r o r h i z a , showing the habit of the stem with Fig. 7-2. U. fibrosa, a magnified view showing several
whorls of branches bearing many small, nodular traps. traps.

84 / Carnivorous Plants of the United States and Canada

 The flowers of several species of Utricularia
exist in two states, and the factors g o v e r n i n g the a p -
pearance of one or the o t h e r are not k n o w n . A typically
opened and expanded flower is described as c h a s m o g a -
mous. T h e second t y p e , which is barely opened f r o m
the bud stage, and in which the two lips are still folded
with only the spur protruding, is referred to as closed,
or cleistogamous. G e n e r a l l y , cleistogamous flowers a p -
pear earlier in the spring.
S o m e t i m e s the presence of these two flower f o r m s
in the same species is c o n f u s i n g to the b e g i n n e r . In
the past, several professional botanists m a d e errors
and sometimes named the two f o r m s as separate species
w h e n there was only one. I n t e r e s t i n g l y , even cleistoga-
mous flowers, which pollinators c a n n o t enter (or so
we think at p r e s e n t ) , produce capsules of g o o d , viable
seed, as do the c h a s m o g a m o u s flowers of the same
species grown indoors, a w a y f r o m potential pollina-
tors. Since we mentioned in C h a p t e r 6 that by design
the typical flower of this family e n c o u r a g e s c r o s s -
pollination, we have a t h o r n y p r o b l e m to solve in dis-
covering h o w unpollinated flowers of m a n y species of
Utricularia (and of some species of Pinguiculal)
produce seed.
In the a u t u m n , aquatic species of Utricularia o f t e n
f o r m a type of h i b e r n a c u l u m (winter b u d ) called a
turion, and this either floats or sinks to the b o t t o m of
the stream or pool. In the spring, the g r e e n , nodular
hibernaculum expands and then grows into n e w stems
and b r a n c h e s . T h e turion originates f r o m the g r o w i n g
tip of a branch or main stem.
can r e f e r e n c e s , Lloyd, 1 9 4 2 . ) T h e small size o f the trap
made early students reluctant to accept the fact that
the m e c h a n i s m of Utricularia could be so complex and
rapid. We will present a very brief r e s u m e of m o d e r n
t h e o r y as to h o w the trap f u n c t i o n s .
N e a r one end of the trap is a small o p e n i n g sur-
rounded by numerous plant hairs that are often
branched and multicellular. T h e opening is guarded by
a larger, u p p e r - h i n g e d veil of plant tissue called a d o o r ,
and this is f u r t h e r supported in function by a smaller
veil of tissue, the v e l u m , which rests below the door
on a thickened threshold. (See d r a w i n g , p. 84.) T h e
door is w e a k l y sealed against casual entrance by a thin
layer of mucilage. T h e r e are minute glands on the sur-
f a c e of the trap, and located on the interior walls are
peculiar quadrifid ( f o u r - p o i n t e d ) glands.
D u r i n g the resting s t a g e , m u c h of the fluid inside
the b u l b o u s trap is slowly a b s o r b e d , p r o b a b l y by the
Fig. 7-3. A greatly magnified single trap of U. gibba.
still

TRAP FUNCTION
T h e r e is such a long, tedious h i s t o r y b e h i n d the
ultimate disclosure of h o w bladderwort traps function
that we will not detail it here. Suffice it to say that it
reads like a w h o ' s w h o of early b o t a n y . ( S e e general

85 / The Bladderworts
quadrifid glands. As a result, the w a t e r pressure o u t -
side the trap exceeds that w i t h i n , and the sides of the
trap appear pinched in, or c o n c a v e . T h e r e has thus d e -
veloped a n e g a t i v e , or suction, pressure within the trap
which is n o w set to be sprung.
T h e trap is b r o u g h t into action w h e n a small aquatic
animal b r u s h e s by o n e of the sensitive trigger hairs
around the o p e n i n g , or w h e n the trap is o t h e r w i s e
severely disturbed. S t i m u l a t i o n of the hairs apparently
releases an electrical action potential that in turn causes
relaxation of the velum and thus frees the larger door
to suddenly flip b a c k into the interior of the trap b e -
Utricularia T r a p Structure a n d F u n c t i o n
8 6 / Carnivorous Plants of the United States and Canada
cause of the suction f o r c e developed in the resting
stage. T h e opening and suction then allows an inrush
of w a t e r along with the hapless p r e y , after which the
door promptly closes. S i n c e the door is hinged only
one w a y , there is no escape.
O v e r a period of fifteen to thirty m i n u t e s , the trap
resets by a b s o r b i n g w a t e r into its interior and again
recreating a n e g a t i v e suction pressure within. A f t e r a
period of several d a y s , the prey is digested as a result
of e n z y m e activity that has b e e n demonstrated in b l a d -
der e x t r a c t s . T h e e n z y m e s m o s t likely originate f r o m
the quadrifid glands. If the prey is small enough for
sufficient negative pressure to build up while it is in
the trap, it is likely that a n o t h e r animal will be trapped
b e f o r e the one that was caught earlier is completely
digested.
T h e speed of the trapping process has b e e n e s t i -
mated at 1 / 4 6 0 of a second and is far too fast to be
slowed by the m o s t advanced s l o w - m o t i o n c i n e p h o -
tographic techniques. W h e n a m a s s of Utricularia is
lifted f r o m the w a t e r , one can o f t e n hear a fine c r a c k -
ling sound as the traps are sprung and air b u b b l e s pop
into t h e m .
G e n e r a l l y , the prey of Utricularia is quite small, as
one would expect considering the size of the trap. M o s t
catches are minute water i n s e c t s , p r o t o z o a n s , tiny c r u s -
taceans, rotifers, etc. The small waterweed Wolffia
(duckweed) has been f o u n d entrapped in larger b l a d -
ders of U. macrorhiza vulgaris, apparently ingested
when the trap was accidentally sprung and not b e c a u s e
the bladderwort had gone vegetarian. T h e b l a d d e r -
worts are capable of ingesting large n u m b e r s of m o s -
quito larvae, w h i c h is of some universal interest to
m a n , since the quantities ingested are apparently l i m -
ited only by the n u m b e r of traps available. S o m e spe-
cies have even ingested small tadpoles.
T h e r e is a trick to h o w the bladder ingests prey
longer than its longest dimension. Needless to s a y , it
is a gradual process. If the tail of a m o s q u i t o larva is
caught first and the animal is thus held f a s t , the flexible
door closes around the protruding b o d y of the larva
and is still capable of effecting a seal tight e n o u g h to
allow the absorption of water inside the bladder. T h e n
the tail part is digested. If a n o t h e r stimulus o c c u r s , the
reset trap is then able to ingest the remainder or at least
an additional length of the b o d y until the w h o l e animal
is ultimately consumed in successive steps. The
process is very roughly analogous to the spasmodic
swallowing efforts of a s n a k e ingesting a large prey.
8 7 / The Bladderworts
The Species
For purposes of simplicity and easy identification,
we will classify the species of Utricularia b r o a d l y by
flower c o l o r , and then s u b c l a s s i f y according to o t h e r
easily o b s e r v a b l e characteristics that are r e a s o n a b l y
consistent. Professional botanists frequently abhor
such a s y s t e m , since it is not " n a t u r a l " ; that is, it does
n o t express real or theorized evolutionary r e l a t i o n -
s h i p s — r e l a t i o n s h i p s t h a t are themselves o f t e n b a s e d
on only the m o s t tenuous and ephemeral evidence.
H o w e v e r , n o n t e c h n i c a l s y s t e m s f o r the identification of
c o m p l e x g e n e r a are o f t e n m o s t useful to the b e g i n n e r ,
w h o m a y later progress to theoretical considerations if
he so desires.
As y o u read the descriptions b e l o w , you m a y feel
hopelessly mired in the great similarity of m a n y of t h e
species. B u t if you h a v e at h a n d a plant to be identified
and are able to m a k e o b s e r v a t i o n s as you r e a d , y o u
will find that the s y s t e m b e c o m e s w o r k a b l e with some
e x p e r i e n c e . T h i s is not a classical outline, but a linear
k e y , w h i c h the reader m u s t follow f r o m b e g i n n i n g t o
end with any plant. For the convenience of the reader,
the m a j o r categories are listed b e l o w . W i t h the plant
in h a n d or in s i g h t , read f r o m the b e g i n n i n g , e l i m i n a t -
ing inappropriate categories until you c o m e to o n e
that fits t h e plant. T h e n r e f e r to the proper page n u m -
ber f o r detailed discussions of the plant or plants in
that c a t e g o r y .
1. Species with w h i t e flowers (p. 8 8 ) .
2 . Species w i t h purple flowers (pp. 8 9 - 9 0 ) .
A. Aquatic.
B. Terrestrial.
3 . Species with m a i n l y yellow flowers (pp. 9 0 - 9 5 ) .
A. Flowers with fimbriate b r a c t s and sepals.
B . T e r r e s t r i a l ; nonfimbriate b r a c t s o r b r a c t e o l e s .
 C. Flower scapes with radial floats.
D. Aquatic plants w h o s e pedicels arch or recurve
w h e n in fruit.
E. Mainly aquatic plants with occasional subterra-
nean branches and stretches of stem with traps al-
ternating with filamentous " l e a v e s . "
F. Plants with threadlike stems tangled in mats and
floating in shallow water.

The paragraphs below are not meant to be read and

absorbed, but to be scanned as you look for character-
istics of a p l a n t in h a n d .

1 . Species with white flowers. Only one American

species, Utricularia olivacea Wright ex Griseb, regu-
larly h a s w h i t e f l o w e r s . T h i s a q u a t i c p l a n t i s c o n s i d e r e d
by m a n y to be the smallest by weight of any flower-
ing plant in the w o r l d . It has e x t r e m e l y slender, t h r e a d -
like stems with bladders less than 1 mm across. U.
olivacea floats in warm, mainly acid pond waters
among algae and other species of Utricularia and is
quite inconspicuous w h e n not flowering. T h e species
is occasionally found in ponds in the N e w Jersey Pine
Barrens and in the coastal plain f r o m eastern N o r t h
Carolina into Florida. T h e flower, only 2 mm long and
borne on a 2.0-2.5 cm scape, appears from mid-
S e p t e m b e r t o late O c t o b e r a n d , b e c a u s e o f its s m a l l
size, it c a n be easily o v e r l o o k e d . In fact, recent o b s e r -
vations indicate that U. olivacea is probably more
plentiful and widespread than previously supposed.

Fig. 7-4. A small stand of U. o l i v a c e a , the tiny white flowers

barely visible above the surface of the sand. Compare size
with the pine needles in the picture.

Fig. 7-5. The white flowers of U. o l i v a c e a , our smallest blad-

derwort.

88 / Carnivorous Plants of the United States and Canada

2. Species with purple flowers. We will deal with one
aquatic and two terrestrial species.

A. A q u a t i c . — U t r i c u l a r i a purpurea W a l t , is very c o m -
monly found in suitable acid ponds or slow s t r e a m s
over the entire eastern half of the c o n t i n e n t , particu-
larly in the s o u t h e r n coastal plain. T h e long b r o w n
stems float submerged and give rise to whorls of
b r a n c h e s which bear traps. T h e n e w spring g r o w t h
tip often has a bright red color. M a s s e s of plants are
frequently quite large, and f r o m M a y t o S e p t e m b e r
the purple flowers frequently cover the surface of the
water with their bright b l o o m s . T h e flowers are 1 . 0 -
1.3 cm and are b o r n e singly on scapes w h i c h rise 7 - 1 0
cm above the water. T h e lower lip of" the flower has
two lobes, which are saccate ( p o u c h l i k e ) . T h e part of
the scape at water level has a slight f u s i f o r m (tapered
at each end) swelling a b o u t 5 mm l o n g , w h i c h has
never been satisfactorily explained. It m a y possibly
be an actual or primordial flotation m e c h a n i s m to help
keep the flower upright and out of the w a t e r . A w h i t e -
flowered variant of the species, still with saccate lobes
of the lower lip, has recently b e e n found in a pond in
N e w Hampshire.

B . T e r r e s t r i a l . — W h i l e these two species m a y b e o b - Fig. 7 - 6 . A stand of U. purpurea, the bright purple flowers
served in flooded areas, they are basically terrestrial, distinctive above the surface of the water.

as is indicated by the fact that their vegetative p o r -
tions are mainly subterranean. Utricularia amethystina
St. Hil. (sometimes called U. standleyae) is f o u n d o n l y
in low, moist pinewoods of Florida and is rather rare.
It is characterized by a whorl of m i n u t e , bladelike
leaves around the scape at ground level. T h e r e is no
i n f o r m a t i o n on the usual flowering period, and the
flower is occasionally described as w h i t e , pale y e l l o w ,
or whitish purple, these apparently being v a r i a n t s .
Fig. 7 - 7 . A flower of U. purpurea. The saccate lobes of the
lower lip can be seen clearly.
Utricularia resupinata G r e e n e occurs in very wet or
m u c k y areas in m u d flats and along the edges of lakes
and ponds in s o u t h e a s t e r n C a n a d a and the n o r t h e a s t -
ern U n i t e d S t a t e s , and t h e n skips an area until one
reaches S o u t h C a r o l i n a and Florida. T h e plant f r e -
q u e n t l y g r o w s in a h a l f c e n t i m e t e r or so of w a t e r , but

89/The Bladderworts
w i t h the main stem on the m u c k surface or b e l o w .
T h e r e are n a r r o w , small " l e a v e s " f r e q u e n t l y buried i n
the mud. T h e flower appears f r o m M a y t o S e p t e m b e r
and is b o r n e singly on a very t h i n , 2 . 5 - 1 2 . 0 cm scape.
Using y o u r h a n d lens, you will n o t e that the paired
bracts of the scape are j o i n e d , resulting in a tubular
structure. T h e lower lip of the flower is not s a c c a t e ,
as it is in U. purpurea.
3 . S p e c i e s w i t h m a i n l y yellow f l o w e r s . H e r e i s w h e r e
we enter a very difficult a r e a , since m o s t of the N o r t h
A m e r i c a n species are y e l l o w , and some of t h e m are
quite similar. T h e r e is still active discussion as to
w h e t h e r several of these related species are not a c t u -
ally variants o f the s a m e species. W e will use the m o s t
widely accepted classification in this p r e s e n t a t i o n .
A . Flowers with fimbriate b r a c t s and s e p a l s . — T h e r e
is one species, Utricularia simulans Pilger (sometimes
called U. fimbriata), f o u n d flowering t h r o u g h o u t the
y e a r in l o w , m o i s t pinelands of Florida. T h e r e are f r o m
one to seven 7 - m m flowers on a slender s c a p e , and
the main characteristic is that the b r a c t s and sepals
have fimbriate ( t o o t h e d or f e a t h e r e d ) edges.
B. Terrestrial; nonfimbriate bracts or bracteoles.—
W e h a v e three species. R e m e m b e r that the h a b i t a t s
are s u b j e c t to flood, but the b u l k of the plant is firmly
a n c h o r e d in the g r o u n d , the scapes sprout f r o m ground
level, and all species h a v e the t i n y , grassblade-like
"leaves."
1. Utricularia subulata L. is a r a t h e r ubiquitous little
plant occurring in suitably acid, sandy soils and b o g s
t h r o u g h o u t the eastern third of the c o n t i n e n t , b u t it is
m o s t prevalent in the s o u t h e a s t e r n coastal plain. T h e
flowers n u m b e r f r o m three to s e v e n on a w i r y , zig-
zag, 7 cm s c a p e , and t h e y m e a s u r e barely 8 mm across.
90 / Carnivorous Plants of the United States and Canada
T h e lower lip is far larger than the upper, and the flat-
tened spur is pressed against the b a c k of the lower lip.
T h i s is the smallest yellow terrestrial species, and it
flowers f r o m M a y to N o v e m b e r . Early flowers may be
cleistogamous and pale yellow.
2 and 3. Utricularia cornuta M i c h x . and Utricularia
juncea V a h l are b o t h larger species with prominent
spurs projecting d o w n w a r d at an angle away f r o m
the lower lip of the flower. U. cornuta occurs in acid,
m o i s t , sandy places and b o g s f r o m M i n n e s o t a east to
N o v a S c o t i a , t h e n d o w n the eastern coastal plain, and
w e s t again to east T e x a s . U. juncea is a more southern
species occurring m a i n l y f r o m the N e w Jersey Pine B a r -
rens south into the coastal plain. B o t h flower f r o m
June to September generally, although U. cornuta
tends to flower earlier in a location where b o t h species
occur together. At first g l a n c e , these species resemble
o n e a n o t h e r , but the following differences are diag-
nostic: U. cornuta has a yellow-green scape which
g r o w s up to 30 cm tall and three to five chasmogamous
flowers that m e a s u r e up to 2 cm long with spurs 7.5—
1 3 . 8 mm long. U. juncea frequently has a purple-green
scape averaging 1 5 - 2 0 c m tall. T h e smaller f l o w e r s
m a y be either cleistogamous or c h a s m o g a m o u s and
m e a s u r e a b o u t 1 . 0 - 1 . 5 c m long i n the c h a s m o g a m o u s
s t a t e , with spurs measuring only 0 . 7 - 2 . 4 cm long.
C. Flower scapes w i t h radial floats.—These are aquatic
plants, and there are two species, Utricularia inflata
W a l t , and Utricularia radiata S m a l l . T h e s e plants occur
in acid ponds and slow s t r e a m s or ditches in the eastern
coastal plain and occasionally inland to Indiana. T h e y
f l o w e r f r o m M a y t o N o v e m b e r . T h e striking charac-
teristic of b o t h these species is a flotation device on
the midpoint of the scape, consisting of air-filled a r m s
radiating out on the surface of the water like the spokes
of a rimless wheel. T h e r e are generally f r o m four to
Fig. 7 - 8 . A stand of U. cornuta filling a bay (moist de-
pression) in Brunswick Co., N.C. These were readily seen
from a passing car; if they had not been flowering, one
would hardly have suspected that they were there.

Fig. 7 - 9 . An inflorescence of U. cornuta showing its large,

bright yellow-gold flowers with long vertical spurs.

Fig. 7 - 1 0 . U. subulata, a small terrestrial that is very com-

mon in its range.

Fig. 7 - 1 1 . A stand of U. juncea. Even though the plants

appear to be standing in water, you should note that they
are firmly anchored in the ground and have simply been
flooded. Closely inspecting and lifting a few plants is often
necessary to distinguish terrestrials from aquatics.

Fig. 7 - 1 2 . U. juncea, photographed at about the same dis-

tance as U. cornuta (fig. 7-9). The flowers are smaller and
some are cleistogamous. The lower portion of the scape
is purple.

Fig. 7 - 1 3 . A fimbriate bract on the scape of yellow-flowered

U. simulans.

91 / The Bladderworts
 Fig. 7-14. A mass of U. inflata floating in the water of a
roadside ditch in Columbus Co., N.C.

Fig. 7-15. U. radiata, a species similar to U. inflata but

much smaller and with fewer "spokes" on the float.

Fig. 7-16. A single scape of U. inflata. The radiate flotation

apparatus can be seen clearly.

92 / Carnivorous Plants of the United States and Canada

ten spokes. T h e arms are divided at the e n d s , w h e r e 2. Utricularia geminiscapa B e n j . is o f t e n referred to
they frequently b e a r traps. T h e s e floats support the as a " s m a l l e r e d i t i o n " of U. macrorhiza. T h i s species
flowering part of the scape out of the w a t e r , while the is limited to s o u t h e a s t e r n C a n a d a and the n o r t h e a s t e r n
lower part of the scape is below w a t e r level and c o n - U n i t e d S t a t e s . It has smaller flowers m e a s u r i n g to 0 . 8
tinuous with the vegetative portion of the plant deeper c m . In addition to its small size, the species differs
in the water. T h e flowers n u m b e r f r o m three to seven in having a less-developed spur without a h o o k . U.
on the average, but there can be up to f o u r t e e n in U. geminiscapa flowers f r o m J u n e to S e p t e m b e r .
inflata. W h i l e there are several differences b e t w e e n
these very similar species (some b o t a n i s t s would still
E. M a i n l y aquatic plants with occasional subterra-
consider radiata only a form of U. inflata), the easiest
n e a n b r a n c h e s and stretches of stem with traps and
to use for identification is size: U. inflata has larger
filamentous " l e a v e s " a l t e r n a t i n g . — T h e r e are two c l o s e -
flowers and a much larger flotation a p p a r a t u s , the
ly related species. Both are literally anchored by
float measuring to 25 cm a c r o s s , with five to ten
b r a n c h e s reaching into the b o t t o m s of ponds, and in
s p o k e s ; U. radiata is far smaller, the float m e a s u r i n g
b o t h , trap f o r m a t i o n occurs in periodic stretches along
6 - 8 cm across with four to seven s p o k e s . U. inflata
the s t e m , m a n y alternate areas being without traps.
also has the interesting characteristic of producing
T h e stem s o m e t i m e s creeps along the b o t t o m s o f ponds
tubers at the ends of some aquatic b r a n c h e s . D i f f e r -
and up into s p h a g n u m t u f t s , where it m a y be growing
ences in flower s p u r s , which h a v e been m e n t i o n e d as
on the open surface and appear s o m e w h a t like a low
differential characteristics in the p a s t , are not reliable
m o s s . B o t h species flower f r o m M a y t o S e p t e m b e r ,
here.
and t h e y r a n g e f r o m the n o r t h e a s t e r n q u a d r a n t of the
U n i t e d S t a t e s into s o u t h e a s t e r n C a n a d a , w e s t w a r d t o
D . Aquatic plants w h o s e pedicels arch o r recurve
A l a s k a , and d o w n into b o g s of the Pacific coastal
w h e n in f r u i t . — T h e r e are t w o species.
range.
1. Utricularia macrorhiza Le Conte (sometimes
1. Utricularia intermedia H a y n e has a flower scape
called U. vulgaris, U. australis, or U. macrorhiza ssp.
5 - 2 0 cm tall that bears three to five flowers up to 2
vulgaris) is a r a n k grower and has the largest b l a d d e r s ,
cm. The flowers of U. intermedia characteristically
measuring up to 3 or 4 mm across. V e r y o f t e n the b l a d -
have a cylindrical spur m o r e t h a n half the length of
ders are dark red to b l a c k . T h e aquatic stems are r o -
and positioned acutely b e h i n d the lower lip.
bust and grow several m e t e r s long. T h e species occurs
2. Utricularia ochroleuca R. Hartman has a similarly
f r o m Labrador to A l a s k a , d o w n the w e s t coast into
long scape w i t h t w o to ten flowers m e a s u r i n g a b o u t
b o g s of the Pacific coastal r a n g e , and in suitable scat-
1.5 c m . In c o n t r a s t to U. intermedia, the spur is very
tered locations throughout the prairie states and f r o m
s h o r t , p y r a m i d a l , and vertical.
the northeastern states to southern V i r g i n i a . T h e scapes
can be 60 cm tall, and they b e a r t w e n t y or more 1.5
cm flowers. T h e spur is well developed and h o o k e d ; F. P l a n t s w i t h threadlike stems tangled in m a t s and
the palate has b r o w n lines on it. T h e flowering period floating in shallow w a t e r . — H e r e we will deal with six
is from M a y through September. species in t w o s u b g r o u p s :

93 / The Bladderworts
 1. L o w e r lip about twice as long as upper, spur
a b o u t h a l f the length of the lower l i p . — T h e r e is one
species, Utricularia minor L. T h i s plant can be found
in b o g s in the Pacific coastal r a n g e , in southeastern
C a n a d a , and in the n o r t h e a s t e r n United S t a t e s . T h e
scape is 5 - 1 5 cm tall and bears two to ten flowers that
are a b o u t 1 cm across. T h e small spur is saccate and
only 1 - 2 m m long. T h e plant flowers f r o m M a y t o
September.
2. Lower lip a b o u t as long as upper, the upper lip
not l o b e d . — T h e spur is almost as long as the lower
lip. T h e r e are five species, and the m o r p h o l o g y of the
flowers is essentially very similar in all, the differences
b e t w e e n species m a i n l y centering on sizes and g e o -
graphic r a n g e s .
a. Utricularia gibba L. has the smallest flower of
this last g r o u p , the scape being 3 - 7 cm tall and bearing
one or t w o 0 . 5 - 1 . 0 cm flowers, which can be seen
f r o m J u n e to S e p t e m b e r . T h e range is Pacific coastal
b o g s and p o n d s and the eastern half of the continent.
b. Utricularia fibrosa Walt, is about the largest
species in this last g r o u p . T h e erect scapes are 1 0 - 4 0
cm tall and h a v e up to seven 1 . 5 - 2 . 0 cm flowers. T h e
species g r o w s in the coastal plain f r o m M a s s a c h u s e t t s
to T e x a s , and it flowers f r o m M a y to N o v e m b e r .
c. Utricularia floridana Nash is mainly confined to
Florida and s o u t h e a s t e r n G e o r g i a and is similar to U.
fibrosa e x c e p t that the scape is flexuous instead of erect,
and the flowers n u m b e r f r o m ten to twenty-five but
are o n l y 1 . 2 - 1 . 5 cm wide.

Fig. 7 - 1 7 . Stems of U. intermedia. You can see that the

bladder-bearing branches alternate with foliar areas.

Fig. 7 - 1 8 . A mass of U. intermedia that has crept up on a

mudflat margin of Lake Michigan.

94 / Carnivorous Plants of the United States and Canada

 d. Utricularia foliosa L. resembles U. floridana in
some respects, but it ranges m o r e widely, f r o m Florida
to Louisiana, and has flowers w h i c h m e a s u r e up to 2
cm. O n e very fine distinction requires a good h a n d
lens and necessitates cutting a cross section of the stem
to examine the vascular b u n d l e s . T h i s species has t w o
sets of vascular b u n d l e s , while all other N o r t h A m e r i -
can species have but one circular ring.
e. Utricularia biflora Lam. is very similar to U.
fibrosa and occurs in m u c h the same r a n g e . T h e differ-
ence in a shorter, 5 - 1 2 cm scape in If. biflora, and there
are usually, but not a l w a y s , o n l y two flowers per
s c a p e — h e n c e the species n a m e .

Fig. 7-19. The flower of U. gibba has much the same mor-
phology as that of U. fibrosa, but the whole plant is
smaller in all respects, the very fine threadlike stems often
being mistaken for strands of algae.

Fig. 7 - 2 0 . Flower of U. fibrosa, a large aquatic, with the

upper and lower lips about equal.

Fig. 7 - 2 1 . A single flower of U. fibrosa. Note the fine red

reticulate markings of the palate.

Fig. 7-22. U. biflora. Very similar to U. fibrosa, except that

the flowers are smaller and there are usually a pair on a
short scape.

95 / The Bladderworts
VIII. Growing North American
Carnivorous Plants

M o s t of the carnivorous plants that h a v e been dis- during millenia of evolution. We can never hope to
c u s s e d a r e n o t t o o difficult t o g r o w s u c c e s s f u l l y , g i v e n duplicate nature exactly, and certainly not by taking
a few basic but rather strict requirements. There is a f e w m i n u t e s to stuff a p l a n t into a p o t , t a k i n g it h o m e
clearly an increased interest in things botanical today, to greenhouse or windowsill, and contentedly con-
and there is special horticultural interest in unusual cluding, " W e l l , that's it."
plants. Carnivorous plants n o w appear frequently in T h i s c h a p t e r i s a r r a n g e d i n five s e c t i o n s . T h e first
general houseplant catalogues, on the shelves of will d e a l w i t h s o m e g e n e r a l p r i n c i p l e s a n d d e f i n i t i o n s
nurseries a n d c o m m e r c i a l g r e e n h o u s e s , and in local dis- of the basic elements of the culture of carnivorous
count and grocery stores. The number of dealers plants. Its m a i n p u r p o s e is to be s u r e t h a t we u n d e r -
specializing in c a r n i v o r o u s p l a n t s is s l o w l y b u t defi- stand important terms and required growing condi-
nitely growing. tions, and to lay out a broad overview of h o w N o r t h
W h a t follows is a highly personalized version of A m e r i c a n carnivorous plants can be cultivated. M o s t
my experience in growing carnivorous plants. I have o f t h e i n f o r m a t i o n y o u will n e e d i s i n t h i s s e c t i o n . T h e
s u c c e s s f u l l y c u l t i v a t e d a n d p r o p a g a t e d n e a r l y all t h e s e c o n d p a r t will d e a l w i t h s p e c i f i c g e n e r a a n d c e r t a i n
species in this b o o k , as well as n u m e r o u s foreign c a r - e x c e p t i o n a l s p e c i e s w i t h i n e a c h g e n u s ; w e will f u n n e l
nivorous plants. I h a v e found that in m a n y cases sev- t h e b a s i c p r i n c i p l e s o f t h e first p a r t i n t o t h e d i s c u s -
eral m e t h o d s will w o r k well, w h i l e i n o t h e r i n s t a n c e s sion of each g e n u s , m e n t i o n i n g certain factors requir-
one m u s t a d h e r e to a specific p r o t o c o l . I do n o t p r e t e n d i n g e m p h a s i s i n e a c h c a s e . T h e r e will b e a t h i r d s e c t i o n
that my m e t h o d s are the only useful ones, and the on how you might manage or construct an outdoor
r e a d e r will v e r y likely c o m e a c r o s s s e v e r a l o t h e r m e t h - b o g . A f o u r t h s e c t i o n w i l l list a f e w s p e c i a l i z e d c o m -
o d s c l a i m i n g e q u a l s u c c e s s . Y o u will d o well t o b e w a r e mercial mail order sources for carnivorous plants na-
of brief a n d o v e r l y simplified instructions for a species tive to North America. Finally, we will say a few
inhabiting certain rather narrow ecologic situations. w o r d s a b o u t field c o l l e c t i n g .
A n d b y all m e a n s r e m e m b e r t h a t a n y p l a n t t h a t h a s
not been specifically bred and developed for the n u r - General Principles
s e r y t r a d e — a n d t h i s i n c l u d e s all o u r c a r n i v o r o u s p l a n t s
— i s never going to adapt perfectly in culture, no mat- DORMANCY.—We will start with the plants' rest
ter h o w apparently successful that culture may be. period, which m a y s e e m a n illogical beginning. But
Plants always g r o w best w h e r e they are found wild, more growth failures occur from ignoring a plant's
assuming that m a n or some other force is not destroy- r e q u i r e m e n t s for d o r m a n c y t h a n f r o m a n y t h i n g else.
ing the s u r r o u n d i n g s to w h i c h the plant h a s a d a p t e d N o t u n d e r s t a n d i n g a n d respecting this resting period

9 6 / Carnivorous Plants of the United States and Canada

has been the main reason for m a n y desperate letters
and phone calls to me f r o m s o m e o n e w h o s e c a r n i v o -
rous plant is dying or just seems to be dying.
All temperate plants have a very definite d o r m a n c y
period, some species of Drosera, Pinguicula, and Utri-
cularia to the extent that they f o r m winter hibernacula.
T h e r e are also more subtle d o r m a n c y patterns in which
the plants simply stop growing for some time each
year. D o r m a n c y is an evolved protective response to
seasonal c h a n g e that might otherwise injure an actively
growing plant. G e n e r a l l y , this e n v i r o n m e n t a l c h a n g e
involves a period of decreased daylight, cooling, d r y -
ing, or a limited combination of these conditions. M o s t
dormancy in the temperate zones of N o r t h A m e r i c a
will occur during the winter m o n t h s , although a s e m i -
dormancy may occur in some plants during late s u m -
mer in response to drying, and we will discuss this later.
T h e r e are indications that d o r m a n c y is the result of
the triggering of complex h o r m o n e m e c h a n i s m s in
plants. T h e m e c h a n i s m s are not yet completely clari-
fied and apparently vary with different species. In the
future it may be possible to control d o r m a n c y in c e r -
tain cultivated plants by various chemical and h o r -
mone applications. But right n o w the best thing to do
about d o r m a n c y is to recognize it and help the plant
to ride it out in the artificial conditions of cultivation.
A plant may indicate that it is going into d o r m a n c y
in several w a y s : (1) A winter h i b e r n a c u l u m will f o r m .
S e a r c h for this in the a u t u m n in appropriate species
where the leaves seem to be failing. (2) T h e plant will
simply stop growing after having put out leaves at a
rather brisk pace. (3) Foliage will dry and die b a c k ,
particularly in rhizomatous plants.
W h e n d o r m a n c y b e g i n s , you m u s t decrease w a t e r -
ing to the point where the soil is just d a m p , and cool
the plant by placing it for the winter in the b a s e m e n t ,
outdoors in a moderately protected area such as n e x t
97 /Growing North American Carnivorous Plants
to a building, or even in the refrigerator wrapped in a
plastic or poly b a g . ( D o not freeze.) If you are g r o w -
ing plants under lights, reduce the p h o t o p e r i o d g r a d u -
ally to mimic the daylight period of the s p e c i e s ' n a t i v e
h a b i t a t at that time of year. An a l m a n a c is helpful
here.
If you try to " f o r c e " a d o r m a n t plant i m p r o p e r l y
with too m u c h w a t e r and w a r m t h , especially in a r e -
duced p h o t o p e r i o d , you are inviting rot and the loss of
t h e plant. A species in d o r m a n c y is not actively
metabolizing and is less able to resist attack by h a r m -
ful bacteria and fungi. Even w h e n n u r s e r y m e n claim
to have forced a n y of the b u l b p l a n t s , you will find
that they have actually provided a m a n a g e d and m o d i -
fied d o r m a n c y period first and h a v e then exposed the
plantings to all the conditions n e c e s s a r y for renewed
growth.
W A T E R . — T h i s would seem to be a ridiculous subject
to spend words o n , but it is i m p o r t a n t . T h e r e is w a t e r ,
and then there is w a t e r .
W h e n g r o w i n g c a r n i v o r o u s p l a n t s , one should use
water that is as pure as possible. " P u r e n e s s " here r e -
fers to a low salts c o n t e n t and the lack of n o x i o u s or
toxic w a s t e m a t e r i a l s . If you h a v e access to an a n a l y s i s ,
the total solids c o n t e n t of good w a t e r f o r carnivorous
plants should be less than 50 parts per million (equiva-
lent t o 1 0 0 m i c r o m h o s o f electrical c o n d u c t i v i t y ) . T h e
water should be neutral or acid r a t h e r than hard and
alkaline. Ideally, such w a t e r c o m e s f r o m collections of
r a i n w a t e r , a g r a n i t e b e d r o c k well or spring ( l i m e s t o n e -
b a s e d wells o f t e n yield w a t e r w i t h too m a n y salts in
i t ) , distilled w a t e r , and rarely local tap w a t e r , w h i c h
should ideally stand f o r t w e n t y - f o u r hours to allow
t h e release of any chlorine. T h e r e is a n e w process
and appropriate apparatus, called reverse osmosis,
w h i c h produces pure w a t e r that is almost the e q u i v a -
lent of distilled, and it does so rapidly and cheaply and
with less f u s s . Do not use any of the various water
softeners intended for " p u r i f y i n g " and s o f t e n i n g water
for h o m e use. T h e s e usually e x c h a n g e one set of toxic
chemicals for a n o t h e r , " t o x i c " as far as plants are c o n -
cerned. Y o u m a y use certain m o r e expensive double
i o n - e x c h a n g e resin setups to desalt w a t e r . Be cautious
of using w a t e r f r o m local rivers and c r e e k s , since m o s t
of these w a t e r w a y s are n o w heavily c o n t a m i n a t e d with
toxic industrial and sewage w a s t e s as well as with r u n -
off of fertilizer f r o m f a r m l a n d s .
In g e n e r a l , during the active g r o w i n g s e a s o n , m o s t
terrestial carnivorous plants require e v e n , c o n s t a n t
m o i s t u r e . T h e soil should definitely be wet to the
t o u c h , but not sopping and dripping.
H U M I D I T Y . — T h i s f a c t o r i s very i m p o r t a n t . Since
nearly all our carnivorous plants are native to a bog
of one sort or a n o t h e r , all require a good degree of
humidity to g r o w well. In b o g s and s w a m p s , there is
abundant surface moisture which evaporates and
raises the relative humidity to saturation on w a r m
days w h e n plants are physiologically active. T h e plants
present in a b o g are adapted to such a situation and
h a v e c o m e to require a good relative humidity to r e -
main h e a l t h y . M o s t h o m e s , especially w h e n heated i n
the winter and air conditioned in the s u m m e r , are f a r
too dry to support h e a l t h y g r o w t h if a carnivorous
plant is placed casually on a coffee table or w i n d o w -
sill. H u m i d i t y will o f t e n have to be a u g m e n t e d in arti-
ficial g r o w i n g conditions.
T h e r e are several w a y s to increase humidity for y o u r
plants. O n e is to g r o w t h e m outdoors if you live in the
eastern third of the c o u n t r y or on the Pacific slopes in
the n o r t h w e s t , w h e r e there is a b u n d a n t natural h u -
midity during the w a r m s e a s o n s . Further inland, it is
98 / Carnivorous Plants of the United States and Canada
too dry. An ideal situation is to have access to a green-
h o u s e where humidity can be carefully controlled. A n -
o t h e r is to use a terrarium setup of pots enclosed in a
glass case and lifted above water level. A n d here I will
put in a word of c a u t i o n : Do not under any circum-
stances expose a closed small greenhouse or terrarium
to direct sunlight for any protracted period. In such a
closed c h a m b e r , temperatures can and will rise pre-
cipitously to dangerously high levels, and this will
kill plants in a short time.
A m e t h o d of raising humidity in a room environ-
m e n t is to use one of the several efficient commercial
humidifiers available on the m a r k e t . S u c h equipment
will often raise the humidity to such an extent that fur-
niture will be ruined, so you will have to keep your
plants in a w a t e r - s a f e area. Finally, the old saw about
placing y o u r pots of plants on a tray of moist pebbles
does not really w o r k at all well.
L I G H T . — C o n s i s t e n t with our thesis that plants r e -
spond b e s t to e n v i r o n m e n t a l factors that exist where
they e v o l v e d , sunlight is the best source of light for
carnivorous plants. A minority of species (to be dis-
cussed later) require partial shading at some time
during the g r o w i n g c y c l e , but m o s t do well in full
sunlight. N a t u r a l light has the proper intensity and
spectral composition f o r the best development and
g r o w t h o f plants.
N o t e v e r y o n e has access to a g r e e n h o u s e , and p e r -
haps local climate precludes keeping carnivorous plants
outdoors. T h e increased use of various sources of a r -
tificial light has c o m e a b o u t , m o s t l y with excellent
results. W e r e c o m m e n d the use o f fluorescent lights,
r a t h e r than i n c a n d e s c e n t , b e c a u s e of the problems with
t e m p e r a t u r e and spectral composition that arise with
the latter. Equally effective are cool white fluorescent
tubes and the more expensive but supposedly longer
lasting special growth tubes. The best minimum
fixture is one with a white or aluminized reflector
and four 4 8 - i n c h , 4 0 - w a t t tubes w h i c h produce an
intensely lit central area of usable size. Keep in
mind that there is considerable loss of intensity at
the ends of fluorescent tubes and out laterally f r o m
the fixture. T h e photoperiod is adjustable with avail-
able inexpensive automatic timers, so the grower m a y
exactly duplicate seasonal c h a n g e s . Fluorescent lights
are cool, but they can be made even more so by r e -
m o v i n g the balast f r o m the fixture, l e n g t h e n i n g the
wires, and then mounting the balast a w a y f r o m the
growing area. O f t e n , the m i n i m a l w a r m t h of fluor-
escent light fixtures is desirable w h e n g r o w i n g plants
in unheated b a s e m e n t s . Humidity can be maintained
with small tabletop humidifiers or with terrarium set-
ups, which will be quite safe since, with fluorescent
light, internal temperature buildup will not be a factor.
I have had considerable experience with fluorescent
lights and still use t h e m for handling certain seeds and
cuttings that require precise a d j u s t m e n t s of light i n -
t e n s i t y , photoperiod, t e m p e r a t u r e , and humidity. T h e r e
is one m a j o r problem with the use of artificial l i g h t s ,
and this involves the drop-off of light intensity as lights
are raised farther a w a y f r o m plant surfaces. T h e i n -
tensity of light varies inversely with the square of the
distance of the source f r o m the plant surfaces. To il-
lustrate, if lights are set 20 cm f r o m the plant surface
and the grower then moves them up to 40 cm (twice
the d i s t a n c e ) , the light intensity is not cut in h a l f , but
cut down to o n e - f o u r t h . T h i s is no p r o b l e m w h e n
growing prostrate rosette f o r m s such as Dionaea, m o s t
species of Drosera, Pinguicula, and some species of
Sarracenia, because you can set the light at the optimal
distance and leave it. But there is a p r o b l e m with
9 9 / G r o w i n g North American Carnivorous Plants
growing the tall, erect p i t c h e r plants and with m a i n -
taining flowers on taller s c a p e s ; b o t h require lifting the
light fixtures to a greater height as the plants g r o w .
As o n e raises the light source to a c c o m m o d a t e a rapid-
ly g r o w i n g erect Sarracenia pitcher, o n e is decreasing
the light i n t e n s i t y to the g r o w t h c r o w n near the soil
surface w h e r e o t h e r buds will arise. T h e relative d e -
ficiency of light to the n e w leaf buds then results in
e t i o l a t i o n — t h e production of l o n g , leggy, soft green
g r o w t h that falls over easily. T h e s e taller, leggier
pitchers require even more raising of the light b a n k ,
f u r t h e r decreasing light intensity to n e w e r b u d s , and
the spiral c o n t i n u e s until c r o w n rot develops. C o m -
pounding the p r o b l e m is the effect of the decreasing
light o n p r o s t r a t e rosettes that the g r o w e r m a y have
mixed in w i t h the taller species.
T h e o b v i o u s solution is to h a v e several light setups
according to plant h a b i t u s and ultimate height. V e r y
prostrate rosettes (Dionaea, most species of Drosera,
S. psittacina, Pinguicula) u n d e r one setup, intermedi-
ates (S. purpurea, Drosera filiformis typica, etc.) under
a second s e t u p , and the tall, erect p i t c h e r p l a n t s in
some o t h e r sort of a r r a n g e m e n t . I h a v e never f o u n d a
totally s a t i s f a c t o r y solution to the p r o b l e m of raising
the tall species of Sarracenia under l i g h t s , b u t there is
a partial, albeit expensive o n e . T h a t is to use three light
f i x t u r e s — o n e placed a b o v e and one placed at each side
at an angle so that the light is directed toward the soil
surface.
T E M P E R A T U R E . — M o s t p l a n t s respond b e s t t o t e m -
perature variation in a t w e n t y - f o u r h o u r period. Let the
temperature drop 5 ° - 1 0 ° C a t night. T e m p e r a t u r e and
humidity are i n t i m a t e l y c o r r e l a t e d , and light is an
i m p o r t a n t indirect third part of this equation. Increased
light frequently m e a n s rising temperatures and t h e r e -
fore higher humidity r e q u i r e m e n t s . For e x a m p l e , well-
lit plants kept in high relative humidity can tolerate
w a r m e r temperatures than poorly lit or drying plants.
Likewise, good lighting and h i g h e r temperatures are
going to require a higher input of humidity.
1 am o f t e n asked a b o u t the m a x i m u m temperatures
a plant or group of p l a n t s can survive without h a r m .
A g a i n , c o m m o n sense refers us to the native e n v i r o n -
m e n t , and again, one will find a good almanac very
useful. D u r i n g g r o w t h periods it would be nice to
have a daily temperature of 3 0 ° - 3 5 ° C with the opti-
m u m 5 ° C drop at night, but we are seldom so f o r t u -
n a t e . G e n e r a l l y , plants that are native to the n o r t h e r n
reaches are best kept at a temperature rarely exceeding
3 0 ° - 3 2 ° C ; plants indigenous t o the southeast m a y
endure temperatures up to 3 5 ° - 3 7 ° C f o r short periods
during the day. Even wider adaptation can occur in
some cases if the c h a n g e is b r o u g h t a b o u t slowly.
O n e is m o r e likely to find an adaptive plant a m o n g
a b a t c h of seedlings that in m a t u r e plants plucked
f r o m the wild, since in sexual reproduction a r e c o m b i -
nation o f genetic factors occurs. O n e o f these r e c o m -
binations m a y be m o r e adaptive to the n e w artificial
e n v i r o n m e n t , w h e r e a s it would h a v e been selected
against in the original h a b i t a t .
W h a t a b o u t m i n i m u m t e m p e r a t u r e s ? Here again,
we can refer to the situation in nature. But my o w n
experience has disclosed m u c h more adaptability with
lower temperatures than with higher. I have success-
fully g r o w n m o s t of the carnivorous plants discussed
in this b o o k outdoors in central N o r t h C a r o l i n a , where
temperatures reach d o w n t o - 1 8 ° C for short periods
of time on several n o n c o n s e c u t i v e winter nights a year.
An i n t e n s e , prolonged cold snap would be different. (I
recall m a n y an O h i o w i n t e r w h e r e , for days on e n d ,
the high f o r the day was f r e e z i n g — 0 ° C ! ) T h e r e are
confirmed reports of h e a l t h y plants of southern Sarra-
1 0 0 / Carnivorous Plants of the United States and Canada
cenia flava and Dionaea surviving, reproducing, and
of S. flava even hybridizing with native S. purpurea in
P e n n s y l v a n i a b o g s year after year. T h e r e are uncon-
firmed reports of y e a r - t o - y e a r survival of the same
plants outdoors in an even more northern latitude in
M i c h i g a n . B o t h these areas support a population of
native carnivorous plants which would be expected to
survive in local culture.
My conclusion would be to use these examples as
guidelines. N o r t h e r n outdoor growers should protect
plants of southern origin during winters until experi-
m e n t s with one or two expendable plants prove other-
wise. U s e of the tub method of growing allows the
removal of plants to protected areas over w i n t e r ; and
b u r y i n g tubs and pots to the rim will often protect
against severe root freezing. Covering growth crowns
with a mulch over winter can also be helpful, but be
sure to remove the mulch in the spring. S n o w , by the
w a y , is an excellent mulch.
P O T T I N G . — I f you are growing y o u r plants in some
sort of potting rather than in natural or homemade
outdoor b o g s , you will be c o n f r o n t e d with two de-
c i s i o n s : clay or plastic p o t s ; drained or undrained ones.
For a long while it was thought that clay pots were
inherently toxic to carnivorous plants, but now it has
b e e n s h o w n that this is not so. T o x i c i t y was due to the
buildup of a b s o r b e d salts in the walls of older clay
pots, especially p o t s that had previously been used
f o r heavily fertilized plants. Salt buildup in a clay pot
is disclosed by variegated, crusty rings of crystal m a -
terial that do not wash off easily. Y o u can use new
p o t s , but eventually even the minimal quantities of
salts in the soils of carnivorous plants will a b s o r b
and build up in the walls of the clay pots. At this point
you would have to discard the pots, or soak and care-
fully clean each o n e — e x p e n s i v e and tedious processes.
For these reasons I suggest the use of plastic pots and
tubs, with one or two e x c e p t i o n s to be n o t e d later.
Plastic pots do not a b s o r b salts, and their s u r f a c e s are
easily cleaned with good b r u s h e s . No soaking is re-
quired, except to r e m o v e unsightly but h a r m l e s s stains,
if desired.
G e n e r a l l y , it is b e t t e r f o r the b e g i n n e r to use pots
or tubs with drain holes. M o r e f r e q u e n t watering will
be required, but considerable experience and j u d g -
m e n t are needed to use successfully the ultimately
m o r e carefree drainless s y s t e m . Improperly m a n a g e d
pots with no drainage can result in rot and the e v e n -
tual buildup of small a m o u n t s of salts in solution. I
grow most of my specimens of Sarracenia without
drainage. T h i s requires that every year or t w o I spend
some time filling the tubs to the b r i m with w a t e r , then
tipping and e m p t y i n g the excess w a t e r , repeating the
process several times with each tub in order to dilute
and wash out most of the salt buildup. I use drainage
with m o s t other terrestrial plants. O n e should also be
cautious of using a p o t - i n - a - s a u c e r - o f - w a t e r setup;
this a m o u n t s to an undrained s y s t e m unless one r e -
m e m b e r s to flush water t h r o u g h the pots periodically.
If you do use undrained larger pots and t u b s , y o u
will find it helpful to plunge into the g r o w i n g m e d i u m
a smaller pot with drainage holes. T h e drain holes will
allow water to rise up into this smaller pot, and y o u
can use it as a m o n i t o r to c h e c k w a t e r levels. It is also
useful to water your t u b s t h r o u g h this smaller signal
pot. M o s t of the smaller rosette plants do poorly if
w a t e r is applied directly and regularly to their foliage.
Indeed, s p h a g n u m itself can turn b r o w n and die in the
area where even the purest water is applied daily.
W a t e r added to the tub t h r o u g h the signal pot will
settle into the growing m e d i u m f r o m b e n e a t h and
will percolate out sideways and upwards t o w a r d s the
bases of the plants.
1 0 1 / Growing North American Carnivorous Plants
S O I L S . — T h e soils for carnivorous plants should be
loose, p o r o u s , poor in salts and n u t r i e n t s , and h i g h l y
a c i d — w i t h one exception to be noted later. In my e x -
perience, live green sphagnum moss is the best
g r o w i n g m e d i u m f o r the culture of carnivorous plants.
T h e r e are several reasons f o r success w i t h s p h a g -
n u m . M a n y c a r n i v o r o u s plants grow i n s p h a g n u m
b o g s in the first place. Living and growing s p h a g n u m
naturally m a i n t a i n s acidity and a low level of salts
and n u t r i e n t s . S p h a g n u m has unique water-retaining
properties and will m a i n t a i n a proper level of moisture.
At the s a m e t i m e , it is p o r o u s and allows rapid drain-
off of excess w a t e r . S p h a g n u m has natural defense
m e c h a n i s m s against f u n g i and some algae. Finally,
s p h a g n u m is a good health i n d i c a t o r ; the m o s s is quite
sensitive to toxic m a t e r i a l s , and if the s p h a g n u m starts
d y i n g , the c a r n i v o r o u s plants will likely follow.
S p h a g n u m m o s s is actually a g e n u s , Sphagnum, of
a b o u t sixty species o f v a r y i n g colors and textures. W e
do not k n o w of any d o c u m e n t a t i o n that the color in-
dicates acid-producing capability, a l t h o u g h the theory
has b e e n offered i n f o r m a l l y . O n l y e x p e r t s o n mosses
are able to distinguish the species by n a m e . For p r a c -
tical horticultural p u r p o s e s , y o u will be interested in
two kinds o f s p h a g n u m distinguishable b y h a b i t u s :
the c o a r s e , rapidly g r o w i n g species w h i c h are quite
useful for tall, r o b u s t plants such as Sarracenia; and
more compact, smaller, slower-growing sphagnums
w h i c h should be used f o r small rosette plants that
would soon b e o v e r w h e l m e d b y the f a s t e r - g r o w i n g
s p h a g n u m s . O n e can chop coarser s p h a g n u m s up and
use this as a m e d i u m f o r seedlings or small p l a n t s , but
eventually a g r o w t h b u d will t a k e h o l d , and the s p h a g -
n u m will resume its rapid g r o w t h , necessitating r e -
potting.
B y the w a y , s p h a g n u m i s the only " s o i l " w e k n o w
of that g r o w s f o r you. As t h e pots and t u b s begin to
b r i m and overflow with g r o w i n g m o s s , y o u can trim
it off and use the t r i m m i n g s f o r m o r e p l a n t i n g s .
U n f o r t u n a t e l y , live s p h a g n u m is o f t e n h a r d to find
in m a n y areas. Larger nurseries or plant shops m a y be
able to help y o u . B u t do not b u y either so-called sheet
m o s s or milled s p h a g n u m . In m o s t i n s t a n c e s , dealers
will h a v e only dried, dead, b r o w n s p h a g n u m in b a g s
or bales. As long as it is not milled or fertilized, y o u
can use this in the b o t t o m layers of y o u r pots and t u b s ,
b u t y o u should still topdress with a layer of live green
s p h a g n u m . Y o u can t h e r e b y c o m p r o m i s e s o m e w h a t
w h e r e live s p h a g n u m is difficult to get b u t the dried,
so-called long fiber sphagnum in bales is readily
available.
My second choice is to use n a t i v e s a n d y coastal plain
soils, especially t h o s e f r o m the A t l a n t i c s o u t h e a s t . Be
sure to collect it east of t h e clay b a n k s b u t n o t too
close t o t h e o c e a n , w h e r e there m a y b e salt c o n t a m i n a -
tion. C h e c k with state agricultural authorities to be
certain t h a t it is permissible to t r a n s p o r t soils f r o m ,
w i t h i n , or into a state f r o m a particular l o c a t i o n , which
m i g h t b e q u a r a n t i n e d b e c a u s e o f the suspicion that
there are o r g a n i s m s of agricultural disease in the soil.
O f c o u r s e , f e w e r people are going t o b e able t o o b -
tain coastal plain soil than live s p h a g n u m . B u t y o u can
m a k e a p r e t t y g o o d s u b s t i t u t e using the f o l l o w i n g
formula:
1 part fine, w a s h e d silica sand ( R e m e m b e r not to
u s e that collected at the s e a s h o r e — i t will be too
salty.)
1 part fine peat ( U s e G e r m a n or C a n a d i a n , in w h i c h
there is no added fertilizer.)
1 part small g r a d e perlite
M i x this well and w e t it d o w n t h o r o u g h l y ; then let
it stand f o r one or t w o w e e k s to a g e , during which time
it will develop proper acidity and a b a l a n c e d m i c r o -
flora.
1 0 2 / Carnivorous Plants of the United States and Canada
T h e least desirable g r o w i n g medium is plain sand.
W h e n using a n y s a n d — e i t h e r plain or in the above
f o r m u l a — a v o i d lake b o t t o m or river sand. T h e s e sands
will h a v e all the pollutants and nutrients that were
present in their w a t e r s , unless you go to the trouble
of a great deal of w a s h i n g . W a s h i n g m a y not be suc-
cessful a n y w a y , since these sands often h a v e clay
particles w h i c h hold salts and toxins tenaciously. Fine
w h i t e silica sand can be purchased by the b a g in h a r d -
w a r e s t o r e s , w h e r e it is sold for s a n d b o x e s and decora-
tive p u r p o s e s . H o w e v e r , it m a y h a v e been collected
near salt w a t e r , and if y o u h a v e a n y doubt about this,
a quick w a s h with running w a t e r will rid white sand
of a n y salts.
F E R T I L I Z A T I O N . — A f t e r stressing the need for g r o w -
ing media free of n u t r i e n t s and salts, this heading
m i g h t s e e m c o n f u s i n g . B u t if you start with a near
zero level of n u t r i e n t s , and y o u k n o w what you add
(if a n y t h i n g ) , then y o u k n o w e x a c t l y where you stand.
W h i l e m o s t c a r n i v o r o u s plants g r o w nicely in s p h a g -
n u m y e a r a f t e r y e a r w i t h o u t prey or fertilization, they
will b e c o m e m o r e r o b u s t and flower m o r e if very light-
ly fertilized. T h e b e s t w a y to avoid the decision of
w h e t h e r to c h a n c e fertilization is to put your plants
outside on fine days so that they can periodically catch
prey n a t u r a l l y . V e r y little actual prey will do. Feeding
with m e a t s is definitely not advisable. It is too easily
o v e r d o n e , and such highly c o n c e n t r a t e d nutrients re-
sult in plant damage and serve as a breeding ground
for fungus infection.
T h e m o s t useful fertilization s c h e d u l e — a n d I would
n o t start o n e u n l e s s y o u feel y o u r plants are doing
p o o r l y b e c a u s e of a lack of n u t r i e n t s — i s a minimal
m o n t h l y feeding during the growing season. U s e a n y
b a l a n c e d fertilizer diluted to about ten times the dilu-
tion suggested by the m a n u f a c t u r e r ' s label for h o u s e -
plants. If the instructions say o n e t e a s p o o n f u l per
quart for houseplants, m a k e it o n e teaspoon per ten
quarts for carnivorous p l a n t s , or smaller total q u a n t i -
ties in proportion so that the volume c a n be easily
handled. I am partial to seaweed fertilizers r a t h e r t h a n
highly refined materials since these mixtures of natural
origin supply m a n y o t h e r essential elements in trace
amounts. T h e m e t h o d of application is a light, fine,
once-over spray using any of the hand sprayers avail-
able. T h e material will be absorbed foliarly. It is not
necessary to pour solutions into the pitchers of Sar-
racenia.
R e m e m b e r , live s p h a g n u m is a good health m o n i t o r .
If the sphagnum dies and you have recently fertilized,
you may have overdone it. A l s o , a g r o w t h of slimy,
blue-green algae on top of the s p h a g n u m indicates too
much p h o s p h a t e , and likely too m u c h total fertilizer
and salts. By the w a y , I have seen plantings in clay
pots in which there was a nice ring of b l u e - g r e e n algae
on top of the s p h a g n u m near the pot edge o n l y , indi-
cating that salts were being leached f r o m the old pot
into the s p h a g n u m next to the wall.
P E S T S . — T h i s refers to the p a t h o g e n variety, not i n -
quisitive visitors, p e t s , and other people's kids w i t h
carelessly probing fingers. Y e s , carnivorous plants are
susceptible to insect p e s t s , and s o m e others. Plants
grown b y reliable nurseries should b e free o f p e s t s ,
but field-collected plants m a y be a n o t h e r m a t t e r .
We have already m e n t i o n e d the pests of Sarracenia
in C h a p t e r 3. T h e first step in ridding pitcher plants of
Exyra is to seek out any adults resting on the inside
pitcher walls just below the lip. R e m o v e t h e m with
long forceps and kill them. N e x t , if you see telltale
signs of larval activity, such as o r a n g e f r a s s in the
pitcher, a dry ring around the pitcher t o p , or a w e b
spun across the pitcher m o u t h , cut off the afflicted
1 0 3 / Growing North American Carnivorous Plants
pitchers at the very b a s e and b u r n these leaves. T h e
consecutively laid eggs h a t c h sequentially, so y o u m a y
b e t r i m m i n g infested pitchers f o r several w e e k s a f t e r
killing a fertile adult f e m a l e m o t h .
The Sarracenia root borer, Papaipema appassionata,
can be diagnosed by an enlarging conical pile of bright
orange droppings collecting at the g r o w t h c r o w n near
the soil s u r f a c e . If you are l u c k y , you can seek out the
larva w i t h n a r r o w f o r c e p s , pull h i m out of his rhizome
t u n n e l , and destroy h i m . If you cannot remove the lar-
v a , y o u can use a m e t h o d r e c o m m e n d e d by J. A. M a z r i -
m a s which involves the instillation of a dilute solution
of m a l a t h i o n into the larval tunnel with a medicine
dropper. D i l u t e the m a l a t h i o n according to the instruc-
tions on the label. Y o u r plant is likely to develop rot
and die if destruction has b e e n too great b e f o r e the
larva is detected and r e m o v e d . Adults are no problem
in Papaipema.
O t h e r likely pests m o r e peculiar to cultivation are
scale and m e a l y b u g s . S c a l e is an insect appearing as
rows of s o f t tan to hard b r o w n " t u r t l e s h e l l s " a v e r a g -
ing 2 - 4 m m across. T h e y are o f t e n o n the surfaces o f
the g r o w t h c r o w n w h e r e they m a y not be seen and
will o f t e n s h o w up indirectly as deposits of b l a c k
mildew on the upper pitchers. T h i s is because the scale
releases plant juices w h i c h in turn support mildew
g r o w t h . M e a l y b u g s are fluffy, w h i t e , powdery insect
colonies located on leaves. B o t h these insects can in-
fest the interior of Darlingtonia pitchers, where they
are very hard to diagnose and eradicate. T h e t r e a t m e n t
is a light but complete spraying with malathion diluted
according to m a n u f a c t u r e r ' s instructions. T h e scales
will not drop off w h e n they die and dry u p , b u t t h e y
are then h a r m l e s s . T w o w e e k s after the first s p r a y i n g ,
repeat the treatment to deal with any recent egg
h a t c h i n g s . M a l a t h i o n does not affect e g g s , o n l y the
insects. Inspect all n e w plants entering y o u r collection
f o r these pests. T h e y will spread rapidly and hide in
all n o o k s and crannies of the plant's a n a t o m y .
Fungi can be a p r o b l e m with Pinguicula and e s -
pecially with the terrestrial species of Utricularia, less
often with Drosera and Dionaea, and rarely with
Sarracenia. Fungus infections usually indicate that
something is w r o n g with the c u l t u r e : d o r m a n c y r e -
quirements are not being met p r o p e r l y , or s o m e t h i n g is
w r o n g with the proportions of light, w a t e r , and t e m -
perature. Fungus infection is indicated by the m o i s t ,
slimy browning of leaves with rotting of g r o w t h
c r o w n s , w h i c h tends to spread across the pots and
t u b s like a small plague. Y o u m a y also see typical
fuzzy g r o w t h s of fungal fruiting bodies. T h e first step
of t r e a t m e n t is to analyze and correct the culture
p r o b l e m that originally led to the f u n g u s i n f e c t i o n .
T h e second is to apply a fungicide. T h e r e is n o w
available a very effective and safe n e w s y s t e m i c f u n g i -
cide called B e n l a t e or B e n o m y l . M a k e up a suspension
according to the instructions on the label and apply
two w a y s : first, give a good spraying to all leaves in
the pot or t u b ; then u n c a p the sprayer and apply the
solution to the g r o w i n g medium as t h o u g h you were
w a t e r i n g . Y o u will be effectively treating the infection
on the leaves as well as in the w h o l e plant s y s t e m i -
cally t h r o u g h the roots.
Finally, I would again remind y o u that the adults
of the Wyeomyia mosquito larva o f t e n found in pitch-
ers of Sarracenia purpurea are h a r m l e s s to people and
plants.
P R O P A G A T I O N . — T h i s is a fitting finale f o r this s e c -
tion on general principles. It has b e e n said, and rightly
so, that you do not actually have a plant until you are
able to propagate it t h r o u g h seed or vegetative m e a n s .
T h e r e is sound b a s i s for this s t a t e m e n t . If the plant is
doing well e n o u g h to reproduce, and the resulting
1 0 4 / Carnivorous Plants of the United States and Canada
seedlings and buddings in turn g r o w to maturity, then
you are doing something right. We will discuss spe-
cific pollination, seed h a r v e s t i n g , and sowing with
each g e n u s . Here we will mention general procedures.
T h e b e s t medium f o r seedlings is finely chopped,
live green s p h a g n u m mixed with an equal amount of
fine, w a s h e d , white silica sand. Plain sand is second
choice. Place s o m e small granite pebbles in the b o t t o m
of a plastic 2 - 4 inch seedling pot with drain holes,
then put in y o u r soil mix and water thoroughly. N e x t ,
s o w the seeds directly on the surface. Do not sow
too t h i c k l y , and do not cover any carnivorous plant
seeds.
As a general rule, seeds that mature in the spring
or very early s u m m e r are ready f o r immediate sowing
and will g e r m i n a t e promptly. S t o r a g e at ordinary room
t e m p e r a t u r e s will result in deterioration. Seeds that
ripen in the fall m u s t usually undergo a period of
damp cold t r e a t m e n t called stratification. T h i s is best
accomplished by placing the seed-sown pot upright
in a poly b a g , sealing it, and refrigerating it for either
the w h o l e w i n t e r ( f o r g r o w t h in a greenhouse) or a
m i n i m u m of six to eight weeks (for g r o w t h by arti-
ficial l i g h t ) .
Immediately a f t e r sowing spring-ripened seeds, and
a f t e r stratifying fall seeds, place all pots in essentially
the s a m e g r o w i n g conditions as adult plants, using
slightly filtered rather than full sunlight. Germination
of N o r t h A m e r i c a n species will take place in two to
f o u r w e e k s . W h e n w a t e r i n g , use a medicine dropper so
as n o t to displace very tiny seeds. W h e n seedlings ap-
p e a r , it is b e s t to administer a dose of B e n o m y l or
B e n l a t e (diluted according to directions on the l a b e l ) ,
using a medicine dropper. T h i s will help forestall, in
m a n y kinds of seedlings, a c o m m o n fungus disease
called " d a m p i n g o f f . "
Seeds of aquatic species of Utricularia can be sown
directly in the water in which they will be g r o w n
(see following s e c t i o n ) ; those of terrestrials, on soil
surfaces. In fact, terrestrial species of Utricularia and
some other carnivorous plants will prove almost weedy
because they self-seed so readily. Needless to s a y , this
is a rather pleasant weediness.
Seeds, even the tiny spring seeds, can be safely
stored. Allow the collected seeds to dry in the air f o r
a day or two in a quiet place where air currents will
not scatter t h e m . T h e n either place the seeds in air-
tight, dry plastic or glass vials or wrap them in squares
of waxed paper. Label them properly and keep t h e m
in the refrigerator. Under refrigeration, even the m o s t
evanescent seeds keep f o r at least three to five y e a r s .
In cases where it can be used, vegetative p r o p a g a -
tion is the quickest w a y to obtain larger plants of
exactly the same characteristics as the parent plants.
T h e r e are several basic techniques that can be used
for N o r t h A m e r i c a n carnivorous p l a n t s :
1. Vegetative apomixis can be used for propagation
w h e n it occurs in Dionaea or Drosera intermedia.
C h a p t e r 2, p. 21.) Carefully remove the plantlets f r o m
the scapes and plant them so they can take root.
2. Natural leaf budding will occur often in Pinguicula
primuliflora, occasionally in other Gulf species
Pinguicula, and fairly c o m m o n l y in m a n y species of
Drosera. W h e n the buds are large enough and have
developed root s y s t e m s of their o w n , they can be
separated and planted.
3. Stolons will send up plants in Darlingtonia,
these can be cut loose and replanted after they develop
their own root systems.
4. Rhizome branchings or buddings of Sarracenia that
occur naturally in successful culture m a y be separated
by cutting after you are sure that roots are being p r o -
duced on the rhizome b r a n c h .
1 0 5 / G r o w i n g North American Carnivorous Plants
5 . Rhizome cutting m a y b e used for the propagation
of Sarracenia. I am grateful to Steve Clemesha of
Australia f o r developing and telling me a b o u t the fol-
lowing technique. U n c o v e r the upper half of a h o r i -
zontal rhizome of a good-sized plant until the top of
the stem is fully e x p o s e d , but leave the roots in the
soil. Using a s h a r p , f r e s h , single-edged razor b l a d e ,
very carefully slice perpendicularly into the rhizome
a b o u t h a l f w a y . R e p e a t this at several points. Leave the
top of the rhizome u n c o v e r e d , and in a f e w w e e k s n e w
g r o w t h buds will appear at the sites of the cuts. As
soon as n e w roots d e v e l o p , you can complete the sepa-
ration of y o u r n e w individual plants.
6. Leaf cuttings w o r k very well with most species of
Drosera, m o d e r a t e l y well with n o n b u d d i n g species of
Pinguicula, and fairly well with Dionaea. The pro-
cedure is b e s t done early in the g r o w i n g season. T h e
technique is to cut off a fairly fresh but m a t u r e leaf
at the base of the petiole and to place the entire leaf
right side up on a b e d of m o i s t , finely chopped green
s p h a g n u m . T h e lower side of the leaf must be flat on
(See the surface of the s p h a g n u m , and you can achieve this
by pinning it with t o o t h p i c k s or by spreading a single
layer of coarse cheesecloth over m o u n d e d s p h a g n u m .
T h e plantlets will c o m e up t h r o u g h the holes in the
of cloth. David Kutt originated this idea. Leaves of
Dionaea and Pinguicula will s o m e t i m e s do b e t t e r if the
end of the petiole is placed in the s p h a g n u m rather
t h a n simply being laid on top. T r y several cuttings
both w a y s .
and N e x t , place y o u r pot of cuttings in a h i g h - h u m i d i t y
c h a m b e r in the shade and in a w a r m — b u t not h o t —
place. T h e floor of the g r e e n h o u s e under a b e n c h is
excellent in the spring. O v e r a period of several w e e k s
you will note the a p p e a r a n c e of plantlets f r o m the
margins and surfaces of the flattened-out leaves and
at the petiole end of some leaves. Let these g r o w until
y o u are certain that roots h a v e f o r m e d . D u r i n g this
period the m o t h e r leaf will usually b l a c k e n and die.
W a t c h out f o r fungal g r o w t h , and treat it appropri-
ately if it appears. W h e n the y o u n g plants are well
rooted, transplant them carefully to separate pots and
slowly acclimate them to proper lighting and decreased
humidity o v e r a period of several m o r e w e e k s .
Genus Notes
N o w that w e h a v e discussed s o m e basic principles
of g r o w i n g carnivorous plants native to N o r t h A m e r i -
c a , we will m e n t i o n each g e n u s and relate i m p o r t a n t
specifics, e m p h a s e s , and e x c e p t i o n s .
D I O N A E A . — D i o n a e a requires a complete period of
d o r m a n c y as outlined in the previous section. W h e n
d o r m a n c y b e g i n s in cultivation, the leaves usually turn
b l a c k and w i t h e r , but the rhizome r e m a i n s h e a l t h y
and fleshy as long as there is no attempt to f o r c e
g r o w t h . U s e a small variety of s p h a g n u m or the coastal
plain soil m i x ; drainage is b e s t . W h e n the plants are
actively g r o w i n g , there should b e a b u n d a n t light, h u -
m i d i t y , and m o d e r a t e b u t w a r m t e m p e r a t u r e s .
Dionaea's spring g r o w t h pattern b e g i n s with a f e w
trap leaves that are not v e r y large or well developed,
followed by the flower scape if the plant is large
e n o u g h to b l o o m . A f t e r flowering, larger and more
typical leaves are produced all season long. M a n y will
not wish to b o t h e r w i t h the flower, and it can be cut
off at an early s t a g e , w h i c h tends to stimulate the f o r -
m a t i o n of larger and earlier traps.
I have found that the b e s t routine propagation
m e t h o d is by seed. Y o u should h a v e two or more plants
in flower s i m u l t a n e o u s l y , since selfing is very difficult
b e c a u s e of the differing m a t u r a t i o n periods of anthers
and stigmas in the s a m e flower. O n e c a n pluck indi-
1 0 6 / Carnivorous Plants of the United States and Canada
vidual stamens with fine forceps and touch these to
t h e stigmas of a n o t h e r plant, but this is tedious. A
quicker and more certain method is simply to bend the
scapes and b r u s h the open faces of two flowers on two
different plants lightly against each other with a cir-
cular m o t i o n . T h i s will result in a mutual transfer of
adequate pollen f r o m one to the other. T h e flowers of
Dionaea open successively, and you will have to repeat
the pollination process daily as n e w flowers open. T h e
small, b l a c k seeds m a t u r e in six weeks and can be
sown immediately as described in the previous section.
S A R R A C E N I A . — T h i s genus also requires very care-
ful attention to d o r m a n c y . If you are growing the
plants in an undrained t u b , r e m e m b e r to cool the tub
and cut w a y b a c k on watering until the sphagnum is
just d a m p . D u r i n g the active growing season, give
plenty of light, w a t e r , and moderate w a r m t h , and grow
the p l a n t s in green s p h a g n u m . D a i l y misting of the
foliage with w a t e r is also beneficial.
W h i l e m o s t species of Sarracenia do well in full
sun, S. purpurea requires a bit of special care. Give it
full sun early in the spring to encourage flowering and
good pitcher f o r m a t i o n . (Flowers can be clipped off in
bud if n o t desired.) T h e n , as the days of summer b e -
c o m e h o t , you should place the plant in partial shade
and protect it f r o m breezes. T h i s species does better
with s o m e w h a t higher humidity allowances than other
m e m b e r s o f the g e n u s .
L e a f cuttings have never b e e n regularly successful
as a propagative m e t h o d f o r Sarracenia. T h e separation
of natural rhizome buds and branchings and the
partial rhizome cutting technique described in the
previous section w o r k very well. Seed is not at all dif-
ficult if one is willing to follow instructions for stratifi-
cation. If the plants are not outdoors, you will have to
pollinate the flowers. A l l o w the flower to mature by
being open three to five days. T h e n lift a petal and
gather a small quantity of pollen f r o m the umbrella
cup with the flat edge of a toothpick. D u s t the pollen
over the small stigma lobes on the inside surface of the
umbrella points. O n l y one lobe need actually be polli-
nated, but do t w o or three to be sure. Y o u can self-
pollinate or cross b e t w e e n plants. Seeds will set by
autumn, the drying and enlarging capsule eventually
splitting. R e m o v e the entire capsule, open it c o m p l e t e -
ly, and separate the seeds f r o m their a t t a c h m e n t s onto
a sheet of paper.
The various species of Sarracenia hybridize very
readily, and you may wish to try y o u r hand with
simultaneous flowers of two different species. R e m e m -
ber to label properly.
D A R L I N G T O N I A . — I have f o u n d this t o b e the m o s t
difficult N o r t h A m e r i c a n carnivorous plant to g r o w
truly successfully. I am not referring to a plant that
seems to barely hang on f o r a year or t w o or t h r e e , b u t
one that year a f t e r year produces n e w , larger, and
more vigorous g r o w t h with active stolon production.
U n f o r t u n a t e l y , this species is being offered m o r e by
general nurseries under the illusion that it is an easy
plant to keep in all areas of the c o u n t r y . S o m e n u r -
series even display the plants, usually newly shipped-
in adult specimens stuck into a pot of s p h a g n u m . As
the display plants s u c c u m b , they are promptly r e -
placed by more f r o m the refrigerator.
A f t e r much trial and too m u c h error, I have f o u n d
the following methods to be m o s t successful for me.
First of all, this is one plant that should be g r o w n in
a clay pot, a new and unsalted o n e , of course. T h e
reason f o r using clay is that water seeps into and
through the pot wall and then e v a p o r a t e s , t h u s cool-
ing the pot and roots. R e m e m b e r , this plant's n a t i v e
h o m e is in some very cold-water w e s t e r n b o g s . T h e
1 0 7 / G r o w i n g North American Carnivorous Plants
growing medium m u s t be particularly loose and very
poor in solubles. Y o u can use a m i x t u r e of live, green
s p h a g n u m o f the coarse variety and very coarse p e r -
l i t e ; or as I p r e f e r , simply a potful of w a s h e d , coarse
granite gravel w i t h live s p h a g n u m as a topdressing.
S u c h a potting will encourage a good supply of o x y g e n
to the r o o t s , as well as maintain humidity and cooling
in the r o o t area in spite of excellent drainage. R i n s e
liberal quantities of cold w a t e r through the pot daily.
K e e p the plant in a cool, h u m i d , semishaded place in
s u m m e r . If you live in areas with very w a r m s u m m e r s ,
and n i g h t t i m e t e m p e r a t u r e s d o not drop below 2 0 ° -
22 °C f o r m a n y n i g h t s in succession, you will very
likely lose y o u r plant. N e e d l e s s to say, a proper period
of d o r m a n c y is required f o r good health.
If y o u r plant is doing well, it will produce stolons
a f t e r a y e a r or t w o , and o n c e the buds are well rooted,
you can cut and separate t h e m as n e w plants. Or if you
are using a large clay p o t , let it fill w i t h a mass of
pitchers.
Seeds are easily produced by selfing or cross-polli-
nating flowers that h a v e b e e n open three to five days.
Lift a p e t a l , w i t h f o r c e p s pluck off a s t a m e n or t w o ,
and b r u s h the anthers over the stellate stigma b e n e a t h
the " b e l l . " S e e d s will set by fall and can be harvested
and germinated as in Sarracenia.
I r e - e m p h a s i z e that this species is not an easy long-
term s u b j e c t , and plants are e x p e n s i v e . B u t it is very
a t t r a c t i v e , and m a n y will wish to try it.
D R O S E R A . — Y o u should b e able t o g r o w and p r o p a -
gate this g e n u s very easily in smaller s p h a g n u m s ,
chopped s p h a g n u m , or coastal plain soil m i x . Do n o t
b e c o m e worried o v e r the species that f o r m hibernacula
in the fall. T h e s e include D. linearis, D. anglica, D. ro-
tundifolia, D. intermedia, and both forms of D. fili-
formis. T h e h i b e r n a c u l a of D. fdiformis v. typica s e e m
especially discouraging at this s t a g e , since they are
well covered with a dense coat of b l a c k hairs t h a t , w h e n
moist, m a k e the w h o l e thing l o o k as though it has
rotted. But careful inspection by separating some of
the hairs discloses the tight bud of bright green pri-
mordial leaves. R e m e m b e r to cool and dry out the
plants to the point that they are barely damp for w i n -
ter. D u r i n g the growing season, give all species very
good light f o r m a x i m u m coloration and development.
D. linearis, as you will recall, will not be readily
available, and it occurs naturally in marl or alkaline
b o g s instead o f acid situations. W e have f o u n d , h o w -
ever, that it adapts to acid s y s t e m s in cultivation. A n -
other good soil formula for this particular species was
developed b y J . A . M a z r i m a s and consists o f equal
parts of vermiculite and fine, white silica sand. Y o u
m a y add some dolomite or not. T h i s mix has m u c h the
same aggregate c o n s i s t e n c y as native marl soils, al-
though it is not alkaline w i t h o u t the dolomite.
P r o p a g a t i o n is by leaf cuttings or seeds. A c t u a l l y ,
y o u will find that m a n y species of Drosera s e l f - s e e d ,
and little plantlets will crop up all over soil surfaces.
If not pollinated by w i n d , insect, or m a n , m o s t Drosera
flowers automatically self-pollinate as they close at the
end of the day. T h e seedpods ripen in f o u r to six
w e e k s , b e c o m i n g plump and d a r k b r o w n t o b l a c k , and
seeds can be collected and sown immediately in m o s t
cases. D. linearis and D. anglica, h o w e v e r , by virtue of
being very n o r t h e r n plants with late seed setting, will
do better with a period of stratification.
P I N G U I C U L A . — O n e m u s t r e m e m b e r that i n their
natural e n v i r o n m e n t s m a n y species of Pinguicula are
shaded over by taller grasses during the hot late s u m -
mer, after having had full sunlight in the cooler
springtime.
1 0 8 / Carnivorous Plants of the United States and Canada
G e n e r a l l y , P. pumila, P. lutea, and P. caerulea need
semishading and drying of the soil to bare dampness
as s u m m e r progresses. D u r i n g the winter, be particu-
larly careful with watering. I have found that placing
the plants on slight m o u n d s in their soils is helpful.
G i v e full light, and w a t e r rather generously in the
spring w h e n n e w leaves and flowers are actively g r o w -
ing. T r y to avoid getting water on the leaves.
T h e G u l f coastal species, P. primuliflora, P. ionan-
tha, and P. planifolia, grow in very wet h a b i t a t s , but
in culture such w e t n e s s predisposes the plants to f u n -
gus attacks during winter d o r m a n c y . T h e r e f o r e , allow
the m e d i u m of even these plants to dry to bare d a m p -
n e s s in the winter. P. primuliflora does well in light
shade as in its native habitat.
P. vulgaris (and the putative "macroceras" subspe-
cies) f o r m winter h i b e r n a c u l a , and you must be espe-
cially careful not to overwater or overheat these. In
f a c t , the b e s t policy is to take the buds u p , dust them
with sulfur, place t h e m in a poly bag with a strand or
t w o of damp s p h a g n u m , and refrigerate them over win-
ter. As you e x a m i n e these winter b u d s , you will often
n o t e smaller offset b u d s , or g e m m a e , at their bases.
T h e s e too will g r o w into smaller, young plants in the
spring.
P. primuliflora will f o r m leafbuds by itself each y e a r ,
and the other G u l f coast species will occasionally. Leaf
cuttings are successful in these species. Y o u can obtain
seeds by pollinating the flowers yourself. O p e n the
flower by grasping the lobes of the upper lip with the
fingers and tear down the lower lip with forceps, thus
exposing the s t a m e n s and stigma. G r a s p a stamen with
y o u r forceps and carefully pull it loose. Y o u will see
the rounded, yellow anther at the tip. R u b the anther
o v e r the o v e r h a n g i n g part of the stigma lobe of the
same or a n o t h e r flower until you see pollen coming
off onto the stigma surface. O v e r a period of four to
six w e e k s , the ovary will swell, turn b r o w n , dry, and
then open to reveal very fine b l a c k seeds. S o w i m -
mediately and treat preventively f o r damp-off.
U T R I C U L A R I A . — T h e terrestrials and s o m e o f the
semiaquatics are no p r o b l e m at all, b e c o m i n g almost
weedy in the pots of other genera. T h e y self-pollinate
and self-seed to the extent that I h a v e followed the
m o v e m e n t of a species f r o m one end of a g r e e n h o u s e
to the other with the air ventilator current. A pinch of
soil in which they are growing can be placed in a n o t h e r
p o t , and it will soon fill with bladderwort plants. S o m e
species, such as U. fibrosa and U. gibba, can be g r o w n
in sphagnum slurries. T h e s e are made by filling a flat
tray half full with live s p h a g n u m and half with water.
T h e obligatory aquatics and semiaquatics you m a y
wish to grow in a pool are a different story. S o m e t i m e s
balanced aquaria w o r k out well, although these m a y
not be acid e n o u g h , and some species of aquarium a n i -
mals eat the Utricularia traps. T h e b e s t w a y to g r o w
these plants is to m a k e a special pool that resembles
their natural habitat. Y o u can either use a large plastic
tub or a child's wading pool sunk in the ground to the
rim, or dig a hole and line it with a continuous sheet of
polyethylene. Place about 5 cm of sandy peat in the
b o t t o m and add pure w a t e r to the t o p , along with a
few strands of live s p h a g n u m and some chips of cedar
w o o d , if available. Let the pool age f o r one w e e k . D u r -
ing this period the wood and peat will settle, and the
water should b e c o m e clear and coffee-colored. If y o u
are familiar with soil-testing k i t s , c h e c k the pH (acid-
ity) of the water. It should be pH 4 - 6 . If the w a t e r is
not acid enough, you can add more s p h a g n u m and peat
and let it age some more. If y o u are familiar w i t h
chemicals, diluted sulfuric acid can be added slowly,
1 0 9 / Growing North American Carnivorous Plants
while stirring, until an initial low pH is r e a c h e d . T h e
p H should n o t b e lower than 4 , h o w e v e r . S u l f u r i c acid,
even w h e n dilute, should b e handled o n l y b y those e x -
perienced in using dangerous chemicals.
N e x t , add y o u r plants or turions. A l l o w for full s u n -
light, and replace the w a t e r as it evaporates. S o m e algae
m a y g r o w at first, b u t you will n o t e that as the p l a n t s
of Utricularia b e c o m e e s t a b l i s h e d , the water will clear,
and it will also seem to buffer at the correct p H .
R e m e m b e r that the aquatics will o f t e n appear to die
off in late s u m m e r or a u t u m n as they f o r m winter buds
or h i b e r n a c u l a . N e w g r o w t h will begin in the spring.
A q u a t i c s are p r o p a g a t e d by simply b r e a k i n g the
stems and placing a piece in a n o t h e r t u b .
An Outdoor Home Bog
If you live east of the Mississippi R i v e r or n o r t h -
w e s t of the Pacific coastal ranges w h e r e there is g o o d
natural h u m i d i t y , you can m a k e artificial outdoor b o g s
of w h a t e v e r size and n u m b e r y o u wish. A f t e r finding
w h i c h plants will adapt, the o n l y limitation will be low
w i n t e r t e m p e r a t u r e s . W i n t e r protection can b e p r o -
vided b y covering the plants. W e have m e n t i o n e d sev-
eral plants endemic to the s o u t h e a s t that have adapted
to winters f a r t h e r n o r t h , and all N o r t h A m e r i c a n c a r -
n i v o r o u s plants should adapt to n o r t h w e s t coastal
regions. Since the r e q u i r e m e n t s of w i n t e r d o r m a n c y
are automatically met in outdoor p l a n t i n g s , m o s t spe-
cies actually do b e t t e r in such a setting than indoors.
First of all, select a good area f o r y o u r bog. It should
receive full sun f o r at least half the d a y , and m o r n i n g
sun is p r e f e r a b l e . For ease of m a i n t e n a n c e , it is b e s t
c o n s t r u c t e d a w a y f r o m deciduous trees and their l e a f -
fall area. T h e r e is no reason w h y a bog c a n n o t be placed
a m o n g other h o m e p l a n t i n g s . Y o u m a y h a v e t o consider
some sort of fencing or o t h e r protection against curious
dogs and cats. Finally, place the b o g c o n v e n i e n t to
w h a t e v e r source of pure water you use. N a t u r a l rain-
fall will do f o r m o s t of the t i m e , but you m a y have to
supplement with w a t e r i n g during s u m m e r dry spells.
N o w for the c o n s t r u c t i o n . Y o u can use o n e or s e v -
eral of the larger plastic tubs available in variety stores,
or you can use any size of the inexpensive children's
plastic wading pools. (Nest three or four of t h e s e ,
since they are thin-walled.) Do n o t plan y o u r bog too
s m a l l ; y o u will always need a larger one t h a n you
originally t h o u g h t . I would t h e r e f o r e strongly suggest
the wading pools. Drill a 0.5 cm hole in the side wall
n e a r the b o t t o m o f w h a t e v e r vessel y o u use. Even
though the o b j e c t of the c o n t a i n e r is to retain w a t e r , a
small drainage hole (which will be b e l o w g r o u n d w h e n
you are done) is helpful to drain off excess water s l o w -
ly after h e a v y rain and to provide a slow flow of f r e s h
w a t e r t h r o u g h the b o g .
N e x t , dig a hole large e n o u g h to a c c o m m o d a t e the
pool or tub at y o u r b o g site. Place the vessel in all the
w a y to its rim and fill in soil a r o u n d the outside to s u p -
port the sidewalls. The surface of your container
should be level. Y o u m a y wish to place natural stone
around the edges to conceal the plastic rim.
N o w y o u m u s t fill y o u r b o g w i t h an appropriate
g r o w i n g m e d i u m . A g a i n I suggest live, green s p h a g -
n u m , p r e f e r a b l y o f the small, t u f t e d , c o m p a c t variety
so that y o u can h a v e m i x e d plantings w i t h o u t being
concerned a b o u t a coarse s p h a g n u m species taking over
the smaller plants. If this t y p e of s p h a g n u m is not
available, use the coastal plains m i x . If live s p h a g n u m
is available in limited quantities and y o u c a n n o t find
enough to fill y o u r b o g , use the coastal plain soil m i x
or dried, baled s p h a g n u m (not milled) in the b o t t o m
and topdress w i t h live s p h a g n u m .
An interesting variation is to sink a smaller tub or

110 / Carnivorous Plants of the United States and Canada

plastic container in your b o g , this smaller one having
no drainage hole. Y o u can prepare this container as a
pond for growing aquatic species of Utricularia
outlined in the previous section.
W a t e r your new bog very t h o r o u g h l y ; this m a y take
a few days if you are using dry s p h a g n u m or soil as
a base. In order to prevent hose water f r o m hitting the
leaves of Drosera and Pinguicula, you may wish
place a small plastic tube or two on the surface of y o u r
b o g , with the tubes connected to a main hose so that
the water will trickle in slowly over the soil surface.
Pinguicula and Drosera do not seem b o t h e r e d by rain
pelting their leaves, as they are by artificial waterings
on a regular basis.
W h e n planting your b o g , try to m a k e the a r r a n g e -
ments look natural. Do not line the plants up in neat
rows but instead use circles, g r o u p i n g s , and curves.
Y o u must follow a broad basic p a t t e r n , h o w e v e r , if you
wish mixed plantings. Place the tall pitcher plants t o -
gether on one half so that they will not shade the other
half of the bog f r o m the m o r n i n g sun. Y o u m a y scat-
ter Sarracenia purpurea among the tall pitchers—they
will get plenty of light for good development early in
the growing season, and later the tall pitchers will
shade and protect them from hot sun and breezes. A l s o ,
plant Pinguicula on the edges of your pitcher plant
stands, raised on little m o u n d s 1 - 2 cm higher than
the soil surface, and ranging in and out a m o n g the
pitcher plants bordering the half of the bog reserved
for smaller plants. T h i s remaining open area can n o w
be used for Drosera, Utricularia, Dionaea, and S.
tacina. A diagrammatic plan for such a b o g is p r e -
sented as an example.
W a t e r your bog when natural rainfall fails and the
tops of the s p h a g n u m tufts seem to be drying. In late
fall, trim off dead foliage (leaving pitcher plant p h y l -
lodia), and if you have severe winters, cover your b o g
1 1 1 / G r o w i n g North American Carnivorous Plants
with black poly sheets pinned down around the edges
with stones. If you leave your bog u n c o v e r e d , r e m o v e
as a n y deciduous tree leaves, particularly o a k , as they
fall on the surface. A large, m o i s t e n e d , flattened d e -
ciduous leaf will smother everything under it, includ-
ing s p h a g n u m m o s s , long b e f o r e the leaf rots. A l s o ,
it has b e e n s h o w n recently that some leaves and o t h e r
to plant p a r t s — a g a i n , particularly those of the o a k — r e -
lease chemical s u b s t a n c e s that inhibit or kill growing
plants of species other than their o w n , a m e c h a n i s m
(allelopathy) that serves to reduce competition in
natural settings and that will just as effectively reduce
y o u r b o g to a s h a m b l e s .
Some Commercial Sources of
Carnivorous Plants
For the c o n v e n i e n c e of readers, I am listing several
nurseries that specialize in carnivorous plants. T h e r e
are m a n y o t h e r dealers that sell o n e or two species, or
that obtain their material f r o m one of the sources b e -
l o w , but I have not included them since they are sec-
o n d a r y and their lists are limited. M a n y dealers in
wild flower plants also have for sale carnivorous spe-
cies that are native to their areas.
I suggest writing several of the dealers to obtain
their catalogues. Do not f o r g e t to enclose m o n e y f o r
the cost of the c a t a l o g u e , if n e c e s s a r y . Look the lists
o v e r carefully and c o m p a r e t h e m : prices and sizes of
inventories vary greatly. Practically all have quality
psit- material and use proper p a c k i n g and shipping p r o -
cedures. Many offer interesting foreign species as
well. T h e best time to order y o u r plants is in very early
spring. If offered, request optional special handling or
air mail delivery.
W h e n y o u r plants arrive, open the p a c k a g e and in-
spect t h e m immediately for q u a n t i t y , quality, sizes,
damage, and disease. Then plant your specimens left. M a n y states have laws intended to protect these
p r o m p t l y . T h i s m e a n s preparing your planting area and other valuable native p l a n t s , b u t the statutes are
when you send off y o u r o r d e r , n o t the day the p a c k - u n e n f o r c e a b l e and are all too o f t e n blatantly ignored.
age arrives. Incidentally, those w h o live close to a n y S m a l l collections f o r serious research purposes are
of these nurseries can visit t h e m in p e r s o n and p u r - e x c e p t i o n s , and m o s t states offer permits for this kind
chase p l a n t s , t h e r e b y reducing delays and possible of collecting. C o m m e r c i a l firms are supposed to p r o p a -
d a m a g e and c o n f u s i o n due to mailing. H o u r s are u s u - gate their s t o c k , b u t m a n y have been k n o w n to collect
ally listed in t h e c a t a l o g u e ; if n o t , inquire. regularly f r o m the wild as orders accumulate. S u c h
Finally, this listing is intended for the c o n v e n i e n c e firms should be b o y c o t t e d .
of the reader, and the a u t h o r does n o t personally e n - Y o u m a y be f o r t u n a t e e n o u g h to come into o w n e r -
dorse or stand by a n y of the firms m e n t i o n e d . ship of a natural b o g , or to o b t a i n permission to use
a b o g on s o m e o n e else's property. T h e bog m a y or
S U N D E W ENVIRONMENTS, P . O . Box 5 0 3 , Kenmore m a y n o t c o n t a i n native carnivorous p l a n t s , but even
Station, Boston, Massachusetts 0 2 2 1 5 . C a t a l o g u e — if n o t , y o u can use it for careful plantings. If you are
50£ going to add n o n - n a t i v e carnivorous plants to your
ARMSTRONG ASSOCIATES, BOX 9 4 H , Kennebunk, Maine b o g , be sure to place t h e m in a clearly m a r k e d - o f f area
04043. Catalogue—25^. along one edge so that they will remain segregated.
A R T H U R E. ALLGROVE, North Wilmington, Massachu- Y o u m a y h a v e to do s o m e w o r k on the bog to p r e -
setts 0 1 8 8 7 . C a t a l o g u e — 2 5 ^ . v e n t or partially reverse eutrophication. Cut out u n -
NORTHROP'S INSECTIVOROUS PLANT FARM, P.O. Box 5, w a n t e d s h r u b and h e r b a c e o u s g r o w t h from the center
Hampstead, North Carolina 2 8 4 4 3 . C a t a l o g u e — 2 5 ^ . and f r o m around the edges if the vegetation appears
P E T E R AND P A M , P . O . B o x 4 4 1 5 , S a n F e r n a n d o , C a l i - to be e n c r o a c h i n g on true b o g plants. M a n y bogs are
fornia 9 1 3 4 2 . Free list. partially drained in an ill-conceived attempt to try to
INSECTIVOROUS PLANT ENVIRONMENTS, 2 6 3 8 1 W h i t m a n m a k e the land agriculturally useful. B l o c k up any such
S t r e e t , H a y w a r d , C a l i f o r n i a 9 4 5 4 4 . Free list. efforts at drainage. C h e c k the uphill slopes around a
PETER PAULS N U R S E R I E S , Darcey Road, Canandaigua, b o g , especially w a t e r inflow areas, to be sure that
New York 14424. Catalogue—25^. sources of c o n t a m i n a t i o n and toxic materials are re-
m o v e d . K e e p traffic in the b o g to a m i n i m u m ; much

Field Collecting t r a m p i n g a b o u t damages i m p o r t a n t bog plants and

tends to create p a t h s that soon b e c o m e new drainage

T h e b a s i c rule should b e : D o not collect f r o m t h e ditches. Ideally, y o u can even build a w a l k w a y around

field. I believe the e x c e p t i o n s to this c o n c e p t are less or across the b o g , using creosote-soaked wood or

t h a n one per cent. T r u e , m a n y o f the plants y o u m a y a b a n d o n e d rail ties, w h i c h will do only minimal h a r m

c o m e across s e e m endless in a particular location or to plants w h e r e the wood actually touches t h e m . S u c h

even in a v e r y large g e o g r a p h i c area. B u t equally true a w a l k w a y will reduce i m m e n s e l y the d a m a g e f r o m

is the cliche that, if e v e r y o n e w h o c a m e t h r o u g h t o o k traffic.

some p l a n t s , there would ultimately be f e w or n o n e Y o u m a y be able to stock your b o g or build up y o u r

112 / Carnivorous Plants of the United States and Canada

private growing collection by taking advantage of of damaging the tender leaves of Drosera and Pingui-
dying or threatened stands of carnivorous plants. M a n y cula. Do n o t pile small tender plants but lay t h e m
a bog on private property is undergoing rapid e u t r o p h i - carefully out in flat, shallow trays.
cation, and perhaps the o w n e r does not wish it p r e - W h e n you arrive at the transplant site with y o u r
served. O b t a i n permission to collect in such cases. p l a n t s , c h e c k t h e m carefully for introduceable disease.
Also, m a n y s a v a n n a h s and b o g s are being drained If infested specimens of Sarracenia are found, trim
and cleared for massive forestry and agriculture, some and b u r n all affected pitchers. Plants infested with Sar-
for the right of way for roads, and others are being racenia root b o r e r are b e s t destroyed. In all p l a n t s ,
flooded in dam projects. If you hear of such activity, cut off d y i n g , d a m a g e d , or diseased leaves. It is also
get in touch with the current o w n e r or project m a n a g e r a good policy to w a s h the roots of larger plants clean
and obtain permission to rescue the plants. in some place w h e r e the w a s h i n g s will not c o n t a m i n a t e
M a n y times you will run across a doomed location the n e w planting site. T h e n , plant as soon as possible.
that has far too m a n y plants f o r y o u r own collec-
tion, or perhaps has o t h e r rare and desirable n o n -
carnivorous species as well. In cases of this sort, get
in touch with local botanical g a r d e n s , especially those
that are particularly interested in native plants. S o m e
areas have native plant societies, and these people are
always happy to go out on a " d i g , " as it is called, and
will rescue and relocate endangered native plants by
the truckload. In N o r t h C a r o l i n a , f o r e x a m p l e , the
N o r t h Carolina S o c i e t y for the Preservation o f W i l d
Flowers has often worked in concert with the N o r t h
Carolina Botanical G a r d e n in C h a p e l Hill in various
digs, relocating plants to the garden, which specializes
in native flora, and to other native gardens in the state
as well. T h i s particular a r r a n g e m e n t w o r k s out very
well. But beware of some botanical g a r d e n s , even large
ones. In spite of their prestige, m a n y simply c a n n o t care
f o r carnivorous plants properly, and m a n y donated
collections have b e e n promptly lost.
T h e best time to m a k e field collections (again, do
not collect except for serious research or for the sake
of the plants) is very early spring w h e n the plants are
just budding, or in the autumn. A l w a y s collect a ball
of the soil or s p h a g n u m in which the plants are g r o w -
ing to help maintain t h e m during transport. Be careful

113 / G r o w i n g North American Carnivorous Plants

Additional Reading
T h e following articles and b o o k s were selected to v o l u m e s , has fairly good coverage of the carnivorous
complement the material in this b o o k . It is not a c o m - p l a n t s , although the i n f o r m a t i o n is brief and obvi-
plete list of writings on carnivorous p l a n t s ; additional ously s o m e w h a t scattered. O n e flaw is gross mis-
i n f o r m a t i o n m a y be obtained f r o m bibliographies in labeling of the Drosera photos. All color photos.
m a n y of the scientific papers, f r o m standard indices, or R o w l a n d , J . T . 1 9 7 5 . C a r n i v o r o u s seedplants: sources
f r o m librarians. S o m e of the entries are a n n o t a t e d as a and references. Hortscience 1 0 : 1 1 2 - 1 4 . A very good
guide. T h i s list is a mixture of popular b o o k s and m a g a - b i b l i o g r a p h y and reading list with m a n y references
zine articles as well as technical materials and journal to consult. A must for serious interest.
papers. T h e n o n t e c h n i c a l material is available t h r o u g h S h e t l e r , S. G . , and M o n t g o m e r y , F. 1 9 6 5 . Insectivorous
any good local l i b r a r y , directly or through interlibrary plants. Leaflet n o . 4 4 7 . Washington, D.C.: Smith-
loan. T h e journal papers m a y be read in university or sonian Institution. T h i s free booklet features good
scientific libraries, or m a y also be obtained through i n - narrative and some fine b l a c k and white photos.
terlibrary loan. S o m e foreign species are mentioned and pictured.
References.
General Z a h l , P. A. 1 9 6 1 . Plants that eat insects. National Geo-
A r g o , V. N. 1 9 6 4 . Insect trapping plants. Natural His- graphic 1 1 9 : 6 4 2 - 5 9 . A very fine color photo article
tory 7 3 : 2 8 - 3 3 . A brief introductory article with good featuring A m e r i c a n species.
photos.
Darwin, C. 1875. Insectivorous plants. New York: D. Venus' Flytrap
A p p l e t o n & C o . A classic w o r k available in m a n y (Dionaea)
libraries in a later A m e r i c a n edition ( 1 8 9 8 ) . A f f o l t e r , J . M . , and O l i v o , R . F . 1 9 7 5 . Action poten-
Lloyd, F. E. 1 9 4 2 . Carnivorous plants. W a l t h a m , M a s s . : tials in V e n u s ' flytraps: L o n g - t e r m observations fol-
C h r o n i c a B o t a n i c a . N o w out o f print b u t possibly lowing capture of prey. American Midland Natural-
republishable, this b o o k provides m u c h i n f o r m a t i o n ist 9 3 : 4 4 3 - 4 5 . T h i s paper and the two that follow
on a n a t o m y , h i s t o l o g y , and early experiments with cover s o m e aspects of electrical potential response in
digestion. T h e r e are m a n y fine line d r a w i n g s , but tissues of Dionaea and the physiology of trap
limited p h o t o s due to w a r t i m e conservation. closure.
Poole, L. and Poole, G. 1 9 6 3 . Insect-eating plants. N e w B e n o l k e n , R. M . , and J a c o b s o n , S. L. 1 9 7 0 . R e s p o n s e
Y o r k : Crowell. A good introductory b o o k for y o u n g properties of a sensory hair excised from V e n u s '
people; line d r a w i n g s . flytrap. Journal of General Physiology 56:64-82.
Rickett, H. W. 1966-1973. Wild flowers of the United J a c o b s o n , S. L. 1 9 6 5 . R e c e p t o r response in V e n u s ' fly-
States. Six regional vols. N e w Y o r k : N e w Y o r k B o - trap. Journal of General Physiology 49:117-29.
tanical G a r d e n . T h i s m o n u m e n t a l w o r k , featuring R o b e r t s , P . R., and O o s t i n g , H . J . 1 9 5 8 . R e s p o n s e s o f
most of our native flowering plants in the regional V e n u s ' flytrap to factors involved in its endemism.

114 / Carnivorous Plants of the United States and Canada

 Ecological Monographs 28:193-218. A very com- . 1920. Another pitcher-plant insect. Entomo-
plete paper discussing m o r p h o l o g y and e c o l o g y ; a logical News 31:90-94.
must for serious interest. . 1 9 2 1 . Pitcher plants and their m o t h s . Natural
Scala, J. et al. 1 9 6 9 . D i g e s t i v e secretion of Dionaea History 21:296-316.
muscipula (Venus' flytrap). Plant Physiology 44: M c D a n i e l , S. 1971. T h e genus Sarracenia. Bulletin of
3 6 7 - 7 1 . A n analysis o f biochemical phases o f prey the Tall Timbers Research Station (Tallahassee,
digestion. Florida) 9 : 1 - 3 6 . D o e s not offer a n y t h i n g m o r e t h a n
the Bell, and Bell and C a s e papers a b o v e , but it is
Pitcher Plants all in o n e place. S o m e t a x o n o m i c discrepancies, b u t
(Sarracenia) good descriptions, range m a p s , and a fine set of line
Bell, C. R. 1 9 4 9 . A c y t o t a x o n o m i c study of the S a r r a - drawings of all species.
ceniaceae of North America. Journal of the Elisha M a n d o s s i a n , A. J. 1 9 6 5 . Plant associates of Sarracenia
Mitchell Scientific Society 6 5 : 1 3 7 - 6 6 . This paper and purpurea in acid and alkaline habitats. Michigan
the two that follow comprise a set of classic papers Botanist 4 : 1 0 7 - 1 4 . T h i s paper and the two that f o l -
that present good descriptions, field and natural h i s - low f o r m an interesting and i n f o r m a t i v e series d e -
tory notes, and several fine b l a c k and white p h o t o s . tailing the a u t h o r ' s studies in M i c h i g a n b o g s .
. 1 9 5 2 . Natural hybrids of the genus Sarracenia. . 1 9 6 6 . Variations in the leaf of Sarracenia pur-
Journal of the Mitchell Society 68:55-80. purea. Michigan Botanist 5:26-35.

Bell, C . R . , and C a s e , F . W . 1 9 5 6 . N a t u r a l hybrids i n . 1 9 6 6 . G e r m i n a t i o n of seeds in Sarracenia pur-

the genus Sarracenia, I I : current notes on distribu- purea. Michigan Botanist 5:67-79.

tion. Journal of the Mitchell Society 72:142-52. P l u m m e r , G . L . 1 9 6 3 . Soils o f the pitcher plant h a b i -
C a s e , F. W. 1 9 5 6 . S o m e M i c h i g a n records of Sarracenia tats in the G e o r g i a coastal plain. Ecology 4 4 : 7 2 7 - 3 4 .
purpurea forma heterophylla. Rhodora 58:203-7. Soil structure and analyses of s o u t h e a s t e r n Sarra-
A detailed account of the f o r m of this species free cenia habitats.
of red pigment. P l u m m e r , G . L., and J a c k s o n , T . H . 1 9 6 3 . Bacterial a c -
J o n e s , F. M. 1 9 0 4 . Pitcher-plant insects. Entomological tivities within the s a r c o p h a g u s of the insectivorous
News 1 5 : 1 4 - 1 7 . T h i s paper and the four that follow plant, Sarracenia flava. American Midland Naturalist

comprise a classic series describing various insects 6 9 : 4 6 2 - 6 9 . M o r e study certainly needs t o b e done
that themselves prey upon or live c o m m e n s a l l y with in this area of pitcher plant p h y s i o l o g y and i n t r a -
Sarracenia. T h e r e are excellent drawings and p h o t o s pitcher ecology.
useful for identification, and the writing is clear and P l u m m e r , G . L., and K e t h l e y , J . B . 1 9 6 4 . Foliar a b s o r p -
i n t e r e s t i n g — m o d e l s for some of today's scientist- tion of a m i n o acids, peptides and o t h e r nutrients by
authors! the pitcher plant, Sarracenia flava. Botanical Gazette

. 1907. Pitcher-plant i n s e c t s — I I . Entomological 1 2 5 : 2 4 5 - 6 0 . First reported use o f modern m e t h o d s

News 18:412-20. f o r following absorption and t r a n s f e r using r a d i o -
. 1908. Pitcher-plant i n s e c t s — I I I . Entomologi- isotopes.
cal News 19:150-56. W h e r r y , E . T . 1 9 2 9 . Acidity relations o f the S a r r a -

115 / Additional Reading

 cenias. Journal of the Washington Academy of . 1 9 5 5 . Evidence for the hybrid origin of Dros-
Science 1 9 : 3 7 9 - 9 0 . In this paper the author first d e - era anglica. Rhodora 57:105-30. The author c o n -
scribes the jonesii pitcher plants of the Carolina vincingly argues that hybridization of D. linearis
mountains as a species. C o n t a i n s interesting i n f o r - and D. rotundifolia, followed by amphiploidy, is the
mation on soil pH studies with respect to Sarracenia. origin of this species.
• . 1 9 7 2 . N o t e s on S a r r a c e n i a subspecies. Casta- . 1960. D r o s e r a c e a e . Journal of the Arnold Ar-
nea 3 7 : 1 4 6 - 4 7 . A f t e r years of c o n t r o v e r s y , this p a - boretum 4 1 : 1 5 6 - 6 3 . A brief summation with a k e y ;
per contains the author's decision to consider Sarra- also includes Dionaea.
cenia jonesii a subspecies of S. rubra. S e e also Bell W y n n e , F. E. 1 9 4 4 . Drosera in eastern N o r t h A m e r i c a .
( 1 9 4 9 ) and M c D a n i e l ( 1 9 7 1 ) . Bulletin of the Torrcy Botanical Club 71:166-74. In
. 1 9 3 3 . T h e geographic relations of Sarracenia spite of its date, this is a very nice s u m m a r y ; well
purpurea. Bartonia (15):l-8. In this paper, W h e r r y w r i t t e n , good descriptions.
m a k e s his first presentation of the concept of two
purpurea subspecies. S e e W h e r r y ( 1 9 7 2 ) for his u p - Butterworts
dated t h o u g h t s . (Pinguicula)
W a l c o t t , M. V. 1935. Illustrations of the North Ameri- C a s p e r , S. J. 1962. On Pinguicula macroceras Link in
can pitcher plants. Washington, D.C.: Smithsonian N o r t h A m e r i c a . Rhodora 6 4 : 2 1 2 - 2 1 . G o o d s u m m a -
Institution. A limited edition folio of w a t e r color tion of this possible n e w species split f r o m the P.
prints of all the species, long out of print but avail- vulgaris taxon.
able in m a n y l i b r a r i e s ; a collector's item. A c t u a l l y , G o d f r e y , R. K . , and Stripling, H. L. 1 9 6 1 . A synopsis
the prints are limited b e c a u s e of the material selected of Pinguicula in the southeastern United S t a t e s . The
for illustration, but they are beautifully executed. American Midland Naturalist 66:395-409. Very
T h i s v o l u m e is m o r e important f o r the descriptive good s u m m a r y paper with superb, complete descrip-
notes by Edgar T. W h e r r y and f o r a fine s u m m a r y t i o n s , k e y s , excellent line drawings, and references.
section (with b i b l i o g r a p h y ) on insect associates by
Frank Morton Jones. Bladderworts
(Utricularia)
Sundews T h e r e is a paucity of good general articles on this
(Drosera) genus in N o r t h A m e r i c a , but we understand that this
S h i n n e r s , L. H. 1 9 6 2 . Drosera (Droseraceae) in the will be corrected in the future. Of the references in the
southeastern United S t a t e s : An interim report. Sida general section, Lloyd has an excellent anatomical dis-
1 : 5 3 - 5 9 . T h i s and the following paper present both cussion with m a n y fine line drawings. T h e photos of
sides of the D. brevifolia controversy, which we Utricularia in the R i c k e t t b o o k s are accurately labeled.
mentioned briefly in C h a p t e r 4.
W o o d , C. E. 1 9 6 6 . On the identity of Drosera brevi- C e s k a , A . , and B e l l , M . A . M . 1 9 7 3 . Utricularia i n the
folia. Journal of the Arnold Arboretum. 47:89-99. Pacific n o r t h w e s t . Madrono 2 2 : 7 4 - 8 4 . A particularly

1 1 6 / Carnivorous Plants of the United States and Canada

 useful review, since five species are completely de-
scribed, and all of these occur in the east as well.
Kondo, K. 1 9 7 2 . A comparison of variability in Utri-
cularia cornuta and Utricularia juncea. American
]ournal of Botany 5 9 : 2 3 - 3 7 . A very thorough c o m -
parative description of these two species.
K o n d o , K. (with additional c o m m e n t a r y by Peter T a y -
lor). 1 9 7 3 . A key for the N o r t h A m e r i c a n species of
Utricularia. Carnivorous Plant Newsletter 2:66-69.
An excellent, updated, e a s y - t o - u s e k e y with difficult
points illustrated.
R e i n e r t , G . W . , and G o d f r e y , R . K . 1 9 6 2 . Reappraisal
of Utricularia inflata and U. radiata. American Jour-
nal of Botany 4 9 : 2 1 3 - 2 0 . A n o t h e r good comparison
of two similar species.

Carnivorous Plant Newsletter

Carnivorous Plant Newsletter is a recently conceived
quarterly publication f o r those w h o h a v e a serious in-
terest in the subject and is intended f o r n o n p r o f e s -
sional as well as professional botanists. C P N features
n e w s , short n o t e s , p h o t o s , and reviews of recent litera-
ture and has a seed and plant e x c h a n g e f o r subscribers.
For additional information write one of the c o - e d i t o r s :
J. A. Mazrimas, 3 2 9 Helen W a y , L i v e r m o r e , Calif.
9 4 5 5 0 , o r D . E . S c h n e l l , R t . 4 , B o x 2 7 5 B , Statesville,
N.C.28677.

117 / Additional Reading

Glossary

Actinomorphic. R a d i a l l y s y m m e t r i c a l . Chasmogamous. A term applied to flowers that open

Active trap. A carnivorous plant trap in which a or e x p a n d fully during anthesis.
m o v e m e n t of plant parts takes place during the trap- Cleistogamous. A term applied to flowers that open
ping process. only partially during anthesis.
Ala. Literally, "wing"; a broad, bladelike expansion Clone. I n b o t a n y , a g r o u p o f p l a n t s t h a t all b e a r t h e
of the axial m a r g i n of a pitcher leaf. same genetic composition, having been borne of
Amphiploidy. A p r o c e s s b y w h i c h a n e w s p e c i e s d e - one plant by repeated asexual reproduction.
velops from a hybrid plant. T h e c h r o m o s o m e n u m - Closing trap. A c a r n i v o r o u s p l a n t t r a p i n w h i c h t w o
ber of the hybrid doubles and the plant is capable of identical trap halves a p p r o x i m a t e and thus incarcer-
m a i n t a i n i n g its c h a r a c t e r i s t i c s d u r i n g s e x u a l r e p r o - ate the plant's prey.
d u c t i o n w i t h like p l a n t s . Column. In the context of pitcher plant leaves, the
A n t h e r . T h e tip p o r t i o n o f a s t a m e n , w h i c h p r o d u c e s s t r u c t u r e s u p p o r t i n g a lid o r h o o d .
pollen. Corolla. A c o l l e c t i v e t e r m f o r all t h e p e t a l s o f a f l o w e r .
Anthesis. T h e p e r i o d i n w h i c h a f l o w e r e x p a n d s a n d / Cross-pollination. The exchange of pollen in sexual
or pollination can take place reproduction b e t w e e n t w o different flowering plants.
A s e x u a l reproduction. A f o r m o f r e p r o d u c t i o n i n v o l v - Cuneate. W e d g e - s h a p e d ( a t e r m a p p l i e d t o l e a v e s ) .
ing only o n e parent plant and thus no e x c h a n g e of Cuticle. A w a t e r - i m p e r m e a b l e , w a x y c o a t i n g o f s o m e
genetic material; e.g., b u d d i n g , cuttings, bulb di- plant surfaces.
visions, etc. Door. In Utricularia, the veil of tissue that closes a
Backcrossing. A r e p r o d u c t i v e c r o s s b e t w e e n a h y b r i d trap opening.
a n d o n e o f its p a r e n t p l a n t s . Ensiform. S w o r d - s h a p e d ( a t e r m a p p l i e d t o l e a v e s ) .
Beard. A c o n f l u e n c e o f p l a n t h a i r s o n t h e p a l a t e o f a Enzyme. A c h e m i c a l s u b s t a n c e that s p e e d s a c h e m i c a l
flower. r e a c t i o n w i t h o u t itself c h a n g i n g or b e c o m i n g a c o m -
Binomial nomenclature. T h e m o d e r n s y s t e m o f b i o l o g i - ponent of the reaction.
cal classification w h e r e b y e a c h living o r g a n i s m b e a r s Family. A closely related group of g e n e r a . A f a m i l y
a two-word name corresponding to its g e n u s a n d m a y h a v e only o n e g e n u s , but classification is at
species. the s a m e level as o t h e r families with t w o or m o r e
Bog. A f r e s h w a t e r , c o n s t a n t l y m o i s t o r w e t a r e a d o m i - genera.
nated by mosses and herbaceous plants. Fenestrations. D e p i g m e n t e d , w i n d o w l i k e a r e a s o f p l a n t
Bract. A s m a l l m o d i f i e d l e a f s t r u c t u r e , w h i c h , i n f l o w - t i s s u e , a l s o k n o w n a s areolae.
ers, is located below the calyx. Filiform. T h r e a d l i k e ( a t e r m a p p l i e d t o l e a v e s ) .
Bracteole. A s m a l l b r a c t . Fimbriate. F e a t h e r y , o r v e r y f i n e l y d i v i d e d .
Calyx. A c o l l e c t i v e t e r m f o r all t h e s e p a l s of a f l o w e r . Flypaper t r a p . A c a r n i v o r o u s p l a n t t r a p i n w h i c h t h e

1 1 8 / Carnivorous Plants of the United States and Canada

 victim is ensnared by s t i c k y , mucilagenous s e c r e - plant m o v e m e n t occurs as an integral p a r t of the
tions. trapping process.
Fusiform. Thickened in the middle but tapering Pedicel. A stalk supporting only a single flower.
smoothly toward each e n d . Peduncle. T h e supporting stalk of o n e o r several
G e m m a e . Buds f o r m e d by vegetative reproduction in flowers.
a small cuplike s t r u c t u r e f r o m which they are shed. Petal. T h e o f t e n c o l o r f u l , f o r m - g i v i n g , leaf like p o r t i o n s
G e n u s . T h e first w o r d or m o r e inclusive portion of a of a flower located a b o v e the calyx.
binomial name (pi. genera). Petiole. A leaf stalk.
G r a s s - s e d g e b o g . A sandy bog dominated by grasses P h o t o s y n t h e s i s . T h e s y n t h e s i s o f sugars f r o m c a r b o n
and sedges with scattered longleaf pines. dioxide and w a t e r by green plants with the p a r t i c i -
H i b e r n a c u l u m . A winter bud f r o m w h i c h plants will pation of c h l o r o p h y l .
arise with the return of proper g r o w i n g conditions Phyllodia. L e a f b l a d e - l i k e structures that are p r o b a b l y
(pi. hibernacula). expanded or widened petioles.
H o o d . A pitcher leaf appendage that usually (or d e - Pistil. T h e f e m a l e reproductive p o r t i o n of a flower, in
rivatively) hangs o v e r the pitcher opening. A l s o which seed will f o r m .
called a lid. Pitfall trap. A carnivorous plant trap into which the
Hybrid. G e n e r a l l y , a plant resulting f r o m a cross b e - prey falls a n d c a n n o t exit.
tween two species. P o l y m o r p h i s m . T h e c o n d i t i o n in w h i c h plants of the
K e e l . A ridge on a pitcher plant trap shaped roughly same species (or subspecific classification) have
like the keel of a b o a t . m u c h variation in f o r m .
Lid. See Hood. P r i m o r d i a . Primitive or undeveloped structures such
M a r l bog. A bog in which the " s o i l " is alkaline marl as those a n t e c e d e n t to leaves.
with calcium c a r b o n a t e . Quadrifid. Having f o u r parts or b r a n c h e s .
M a r s h . A tract of wet land, usually with f r e s h , salt, or R a c e m e . A type of inflorescence in which there is a
brackish water to some depth, dominated by taller central stalk w i t h the flowers attached by pedicels.
grasses and reeds. Reticulate. Netlike.
M o r p h o l o g y . In b o t a n y , the f o r m and n o n m i c r o s c o p i c R h i z o m e . A n e l o n g a t e u n d e r g r o u n d s t e m , w h i c h runs
a n a t o m y of plants. a p p r o x i m a t e l y parallel to the s u r f a c e of the g r o u n d ,
O b o v a t e . S o m e w h a t o v a l ; a term applied to a leaf or f r o m w h i c h b r a n c h i n g s m a y arise.
petal which is attached at the n a r r o w end so that Saccate. Saclike.
the distal end appears broader. S a v a n n a h . In the sense used in this b o o k , a sandy
O v a r y . T h e lowermost portion o f the pistil, i n which bog with short grasses and sedges and widely scat-
eggs develop; the ovary will b e c o m e the seed c a p - tered longleaf pines.
sule a f t e r fertilization. S c a l e . A thin, m e m b r a n o u s , colorless, o f t e n brittle d e -
Palate. A p r o m i n e n c e on the lower lip of a s y m p e t a l - generate leaflike structure.
ous, usually zygmorphic flower. S c a p e . A long, n a k e d (without b r a c t s , b r a c t e o l e s , etc.)
Passive trap. A carnivorous plant trap in w h i c h no flowering stem arising f r o m the g r o u n d , usually s u p -

119/Glossary
 porting o n e flower or a tight cluster of flowers at T h r e s h o l d . In Utricularia, the thickened surface against
the very top. which the edge of the door rests.
S e l f - p o l l i n a t i o n . T h e pollination of a stigma with p o l - T r a p d o o r t r a p . A carnivorous plant trap in which an
len f r o m the s a m e flower. appendage closes o v e r an opening and incarcerates
S e p a l . A flower part situated just b e l o w the petals. the plant's prey.
T h e sepal is usually g r e e n , b u t if the flower is t e c h - T u r i o n . In this b o o k , a h i b e r n a c u l u m , but used mainly
nically without p e t a l s , the sepal m a y a s s u m e the in reference to Utricularia.
f o r m and color of a petal. V e g e t a t i v e apomixis. A f o r m of asexual reproduction
S e s s i l e . Set immediately u p o n a n o t h e r structure w i t h - in which plantlets bud f r o m flower parts, including
out an intervening stalk, as a sessile leaf or gland. sepals, petals, s t a m e n s , and pistil.
S e x u a l reproduction. A f o r m of reproduction in which V e g e t a t i v e reproduction. See A s e x u a l reproduction.
s o m e e x c h a n g e o f genetic material occurs b e t w e e n V e l u m . In Utricularia, a m e m b r a n o u s structure for sec-
two o r g a n i s m s . o n d a r y c l o s u r e ; it rests b e l o w the door on the
S p e c i e s . T h e second word or m o s t specific part of a threshold.
binomial n a m e (pi. species). Z y g o m o r p h i c . Bilaterally s y m m e t r i c a l .
Sphagnum bog. A bog dominated by Sphagnum
mosses.
S p u r . In floral m o r p h o l o g y , an e l o n g a t e , closed a p -
pendage of the corolla of a s y m p e t a l o u s flower.
S t a m e n . T h e male reproductive structure of a flower,
consisting of the a n t h e r and its supportive s t r u c t u r e ,
the filament.
S t i g m a . T h e s t i c k y , p o l l e n - r e c e p t i v e , o f t e n k n o b b y top
portion of the pistil.
S t o l o n . A r u n n e r , or any basal b r a n c h that f o r m s roots
and gives rise to an independent plant.
Stratification. In h o r t i c u l t u r e , the process whereby
seeds are exposed to a period of damp cold b e f o r e
they will g e r m i n a t e .
S t y l e . T h e o f t e n c o l u m n a r structure o f the pistil b e -
t w e e n the stigma and o v a r y .
S w a m p . A f r e s h w a t e r a r e a , with w a t e r to some depth,
dominated b y trees.
S y m p e t a l o u s . H a v i n g fused o r joined petals.
S y n g a m e o n . A specialized evolutionary term referring
to plant populations intermediate b e t w e e n the s p e -
cies level and e x t r e m e variants of the s a m e species.

1 2 0 / Carnivorous Plants of the United States and Canada

Derivations of Scientific Names
Understanding the meanings of latinized binomial
names used in biology is an aid in learning to use them
and in remembering them. We have divided the latin-
ized carnivorous plant names used in this b o o k into
two lists b e l o w : a list of generic n a m e s and a second
list of species, subspecies, and f o r m s . In each case we
have attempted to provide the best definition, distilled
from several botanical dictionaries and our o w n e x t e n -
sive reading. It is easy enough to provide a " t r a n s l a -
t i o n " based on Latin or G r e e k roots, but a biologist is
not compelled to explain w h y he named a plant or
animal as he did, and indeed few have. M a n y original
descriptions are lost in antiquity. C o n s e q u e n t l y , a little
detective work and imagination are required to see the
application of a particular n a m e to a particular plant.
In some cases, we have not the faintest idea w h y a
certain name was chosen, even though we may t r a n s -
Drosera. From the G r e e k , meaning d e w y , referring to
the secretions at the tips of glandular hairs.
Pinguicula. Literally, " l i t t l e fat o n e , " referring to the
greasy a p p e a r a n c e and texture of the leaves of this
genus.
Sarracenia. M a n y early b o t a n i s t s were p h y s i c i a n s , as
was D r . M i c h e l Sarrazin ( 1 6 5 9 - 1 7 3 4 ) , who lived in
Q u e b e c and sent specimens of what is now k n o w n
as Sarracenia purpurea to France. His n a m e is n o w
immortalized in this genus.
Utricularia. Literally, little bag or sac, referring to the
bladderlike traps of this g e n u s .
Species, Subspecies and Forms
alata. W i n g e d , referring to the prominent ala of Sarra-

late it, and we will say so in those instances. O n e other cenia alata.

word of caution: Latin or G r e e k language scholars may amethystina. R e d d i s h - p u r p l e in color, like the gem

take exception to some of the botanical meanings b e - a m e t h y s t , referring to the flower color of this Utri-

low. All we can say is that, in those i n s t a n c e s , latiniza- cularia.

tion has been subjugated to b o t a n i z a t i o n ! anglica. Of England, where Drosera anglica occurs
c o m m o n l y and was originally described.
biflora. Two-flowered. The scape of this Utricularia
Generic Names c o m m o n l y , but not a l w a y s , bears two flowers.
brevifolia. S h o r t - l e a f e d , an apt description for this
Darlingtonia. Named after D a r l i n g t o n ; in this case, Drosera.
D r . William D a r l i n g t o n , a nineteenth century n a t u - caerulea. D a r k blue, here referring to the flower color
ralist from Pennsylvania. of this Pinguicula.
Dionaea. T h i s n a m e must have the most romantic ori- californica. O f C a l i f o r n i a .
gin of all. V e n u s ' m o t h e r was D i o n e , and of course capillaris. Literally, hairlike. T h e e t y m o l o g y is unclear,
V e n u s was goddess of love w h o enthralled and but this possibly refers to the m a n y glandular hairs
beguiled all m e n . T h e botanist w h o came up with on the leaves of Drosera capillaris.
this generic n a m e must truly have been beguiled cornuta. From the Latin, m e a n i n g h o r n e d , referring to
by this little plant. the p r o m i n e n t spur of Utricularia cornuta (cf. mac-

1 2 1 / Derivations of Scientific Names

 roceras and microceras below, which are of G r e e k jonesii. C o m m e m o r a t i v e of J o n e s , in this case Frank
origin). M o r t o n J o n e s , an entomologist who studied insect
fibrosa. Having p r o m i n e n t fibers, likely referring to the associates of pitcher plants.
fibrous mats of stems of Utricularia fibrosa. j u n c e a . R u s h l i k e , p r o b a b l y referring to the appearance
filiformis. T h r e a d l i k e , an apt description of the leaves of groups of preflowering scapes in wet sand over-
of this Drosera. laid with shallow water where this Utricularia grows.
fimbriata. Fringed, describing the margins of bracts leucantha. W h i t e - f l o w e r e d , one of the characteristics
and sepals of Utricularia fimbriata. some would use to separate the populations of this
flava. Y e l l o w , w h i c h could describe either the flower species of Drosera from those of D. brevifolia.
petals or pitcher tops of s o m e f o r m s of Sarracenia leucophylla. W h i t e - l e a f e d , appropriate for this white-
flava (cf. lutea b e l o w , also derived from the Latin topped Sarracenia.
and m e a n i n g y e l l o w ) . linearis. Linear, as is the elongate leaf with parallel
floridana. Of Florida, where Utricularia floridana most sides in this Drosera.
c o m m o n l y , but not exclusively, occurs. lutea. Yellow (cf. flava a b o v e ) , referring to the flower
foliosa. V e r y leafy or full of leaves, p r o b a b l y referring color in this Pinguicula.
to the m a n y p h o t o s y n t h e t i c b r a n c h i n g s of Utricu- macroceras. Literally, having a large or long horn
laria foliosa. (-keros, G r . , for h o r n ; cf. Lat. cornuta a b o v e ) , refer-
geminiscapa. Literally, having twin scapes. I have no ring to the longer spur supposedly characteristic of
idea h o w this was applied to Utricularia geminiscapa these populations of Pinguicula that some would set
unless s o m e o n e m i s t o o k the rather arching pedicels aside f r o m P. vulgaris.
for scapes. m a c r o r h i z a . H a v i n g a long root, but this and other
gibba. S w o l l e n . T h e application to this Utricularia is species of Utricularia are rootless, so the reference
unclear, unless it refers to the palate, which is no must be one incorrectly applied at an earlier time to
m o r e swollen than that of any o t h e r m e m b e r of the the very long stem of this plant.
U. fibrosa group. m i c r o c e r a s . Having a short or small horn (cf. macroce-
heterophylla. H a v i n g a leaf variation, here referring ras a b o v e ) , referring to the unusually short-spurred
to the pigment variation (the lack of red) of this far n o r t h e r n variant of this western species of Pin-
northern Sarracenia purpurea. guicula.
inflata. Inflated or swollen, referring to the air-filled m i n o r . S m a l l e r ; perhaps appropriate for the Utricularia
arms of the flotation structure of this Ultricularia. so n a m e d , but not for some of the populations of
intermedia. O b v i o u s l y i n t e r m e d i a t e , although we do Sarracenia we have seen.
not k n o w with respect to w h a t in this Drosera and muscipula. F l y - c a t c h i n g ; eminently appropriate for the
Utricularia. T h e r e f e r e n c e m a y be to leaf length in V e n u s ' flytrap.
the f o r m e r and flower size in the latter. ochroleuca. Y e l l o w i s h - w h i t e . T h e flower of this yel-
ionantha. V i o l e t - f l o w e r e d , referring to the color of the low Utricularia is s o m e w h a t paler than most.
flower of this Pinguicula or its superficial resem- olivacea. O l i v e l i k e . T h i s application is lost in history.
blance to the flower of a violet. It m a y refer to the prominent double ovary (rare in

1 2 2 / Carnivorous Plants of the United States and Canada

 Utricularia) or possibly even to the swelling, m a t u r - villosa. S o f t l y h a i r y ; referring to the p u b e s c e n c e on the
ing green seed capsule. lower portion of the scape of Pinguicula villosa.
oreophila. M o u n t a i n - l o v i n g , referring to the habitat vulgaris. C o m m o n .
of this Sarracenia in n o r t h e a s t e r n A l a b a m a .
planifolia. Flat-leaved. W h i l e curled on the very edges, Some Prefixes
the large leaves of this Pinguicula, w h e n growing in
full sun, are flatter than those of most other species. bi-. Two.
primuliflora. Primrose-flowered, referring to the super- brevi-. S h o r t o r a b b r e v i a t e d .
ficial resemblance the flower of this Pinguicula has gemini-. Twin.
to primroses. hetero-. Other.
psittacina. Parrotlike. A sideview of the pitcher of this leuc-, l e u c o - . W h i t e .
Sarracenia discloses a good case for this epithet, m a c r o - . L a r g e ; sometimes long.
pumila. D w a r f ; this the smallest southern Pinguicula. micro-. Small.

purpurea. Purple, referring to the deep m a r o o n veins ochro-. Yellow (ochre).

or flower parts of Sarracenia purpurea, and to the oreo-. Mountain.

pale purple flower color of the Utricularia species, plani-. Flat.

radiata. R a y e d or radiate, the a r r a n g e m e n t of the primuli-. Primrose (genus Primula).

rotundi-. R o u n d .
spokes of the flotation structure in this Utricularia.
resupinata. Upside down, or apparently s o , as the
general appearance of the flower of this Utricularia. Some Suffixes
rotundifolia. Round-leafed. Although generally the
- a n t h a . Flower,
blade is more o b o v a t e than round in this Drosera,
-ceras. Horn,
the epithet is appropriate,
-flora. Flower,
rubra. R e d . Applicable to pitcher or flower color in
- f o l i a . Leaf,
this Sarracenia.
-leuca. W h i t e .
simulans. Similar to or r e s e m b l i n g ; a relative term
-phila. Loving, or affiliated with,
possibly referring to this Utricularia's resemblance
- p h y l l a . Leaf,
to U. fimbriata.
-rhiza. R o o t ,
standleyae. C o m m e m o r a t i v e of S t a n d l e y ; in this case,
-scapa. Scape.
Paul E. Standley, a noted systematic tropical botanist
of the early part of this century,
subulata. Awl-shaped. The exact reference is not
k n o w n but is likely related to the shape of the flower
spur,
typica. Typical,
venosa. V e i n e d .

1 2 3 /Derivations of Scientific Names

Index
Boldface figures indicate illustrations. annua. See D. brevifolia
Bogs 1-2, 6-9, 8 brevifolia 69-70, 69
Bladderwort. See Utricularia taxonomic problems 69-70
grass-sedge (savannah) 1, 8-9, 8 capillaris 67-68, 70, 6, 68
homemade 109-11 cultivation 107-8
marl 6, 7, difficult identifications 70
sphagnum 7, 9, 59 filiformis 57, 64-67, 70, 99, 107
Butterwort. See Pinguicula v. filiformis (typica) 64, 65, 70, 99, 107, 66
v. tracyi 57, 64, 65, 65
Carnivorous vs. insectivorous 4 flower 56-57
Carnivory, general nature of 1-6 hybrids 70
Conservation 9-10, 54-55, 112-13 intermedia 56, 63-64, 70, 105, 63, 64
Cultivation of carnivorous plants 96-111 leucantha. See D. brevifolia
bog, homemade 109-11 linearis 7, 59-61, 62, 107, 108, 60
dormancy 96-97 longifolia. See D. anglica
fertilization 102-3 rotuniiifolia 56, 58-59, 61, 62, 70, 107, 58, 59
humidity 98 trap 6, 56
light 98-99
fluorescent 98-99 Exyra 27-28, 103, 27
pests 103-4
pots 100-1
Field collections 112-13
propagation 21, 104-6
Flower structure, general 10-12
soils 101-2
temperature 99-100
Insectivorous vs. carnivorous 4
water 97-98
Isodontia 27
Darlingtonia californica 5, 9, 52-55, 103, 105, 107, 53, 55
cultivation 107 Papaipema appassionata 103, 113
flower 53-54, 55 Pinguicula 6, 7, 9, 12, 71-82, 85, 97, 99, 104, 105, 108-9, 111, 113,
habitat 54-55 caerulea 78, 82, 108, 78
historical note 54 cultivation 108-9
insect associates 54 difficult identifications 82
trap 5, 52-53 flower 71-72
Dionaea muscipula 5, 8, 12, 16-21, 73, 99, 100, 104, 105, 106, ionantha 80-81, 108, 81
111, 16, 17 lutea 77, 82, 108, 77
cultivation 106 macroceras 74-75, 82, 108
flower 16, 19 v. macroceras 74-75, 75
habitat 16, 21 v. microceras 74-75
trap 5, 16, 17-20, 15, 17 v. nortensis (nom. nud.) 74-75
Drosera 6, 7, 9, 12, 56-70, 73, 97, 99, 104, 105, 107-8, 111, 113, planifolia 78-79, 82, 108, 73, 78, 79
6, 58, 59 primuliflora 79-80, 82, 105, 108, 80
anglica 61-62, 63, 64, 70, 107, 108, 62 pumila 76, 82, 108, 76

1 2 4 / Carnivorous Plants of the United States and Canada

 trap, 6, 71, 7 2 - 7 3 Sources of carnivorous plants, commercial 1 1 1 - 1 2
villosa 8 1 - 8 2 Sundew. See Drosera
vulgaris 7 4 - 7 5 , 82, 108, 71, 75
Taxonomy, general outline of 1 3 - 1 5
taxonomic problem 7 4 - 7 5
Traps 2, 4 - 6
Pitcher plant, eastern. See Sarracenia
adhesive 6, 6
western. See Darlingtonia
closing 5, 5
Plant names. See Taxonomy pitfall 5, 6
Propagation 21, 1 0 4 - 6 trapdoor 5, 5
leaf budding 105
leaf cuttings 1 0 5 - 6 Utricularia 5, 7, 9, 10, 11, 12, 8 3 - 9 5 , 97, 1 0 4 - 5 , 109, 111, 5
rhizome branchings 105 amethystina 89
rhizome cuttings 105 australis. See U. macrorhiza
seed 1 0 4 - 5 biflora 95, 95
stolons 105 cornuta 90, 91
cultivation 109
vegetative apomixis 21, 105 fibrosa 94, 95, 109, 84, 95
fimbriata. See U. simulans
Sarcophaga 27 floridana 94
flower 8 4 - 8 5
Sarracenia 5, 12, 21, 2 2 - 5 1 , 52, 54, 73, 99, 101, 103, 104, 105,
foliosa 95
106-7, 111, 113
geminiscapa 93
"alabamensis" 46
gibba 94, 109, 5, 85, 95
alata 3 4 - 3 5 , 50, 51, 34, 35
inflata 9 0 - 9 3 , 92
cultivation 1 0 6 - 7
intermedia 93, 94
difficult identifications 51
juncea 90, 91
flava 25, 26, 3 1 - 3 3 , 34, 36, 37, 43, 46, 50, 51, 100, 8, 23, 25,
macrorhiza 87, 93, 84
26, 27, 32, 33
minor 94
polymorphic variants 32, 32, 33
ochroleuca 93
flower 2 4 - 2 5 , 25
olivacea 88, 88
hybrids 4 7 - 5 1 , 48
purpurea 89, 90, 89
insect associates 2 7 - 2 8 , 103
radiata 9 0 - 9 3 , 92
isolation factors 4 9 - 5 0
resupinata 8 9 - 9 0
leucophylla 25, 38, 4 2 - 4 3 , 50, 42, 43
simulans 90, 91
minor 3 7 - 3 9 , 40, 50, 52, 39
oreophila 3 6 - 3 7 , 46, 51, 23, 36 standleyae. See U. amethystina
psittacina 38, 4 0 - 4 1 , 99, 111, 40, 41 subulata 90, 91
purpurea ssp. purpurea 7, 26, 2 8 - 3 1 , 60, 99, 100, 104, 106, 6, trap 5, 8 3 - 8 4 , 8 5 - 8 7
29, 30 vulgaris. See U. macrorhiza
f. heterophylla 28, 2 9 - 3 0 , 30
purpurea ssp. venosa 24, 2 8 - 3 1 , 50, 111, 23, 29, 30 Venus' flytrap. See Dionaea
rubra 37, 4 4 - 4 7 , 45 Vegetative apomixis 21, 105
ssp. jonesii 44, 46, 47, 45
taxonomic problems 4 6 - 4 7 Wyeomyia 30-31, 104
trap 5, 2 2 - 2 4

125 /index
 This book is set in 10 on 13 Palatino
Book design by Virginia Ingram
Photographs by the author
Drawings and maps by Bruce Tucker
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V e n u s ' Flytrap (Dionaea muscipula) Sundews (Drosera)

Eastern North American Pitcher Plants (Sarracenia) Butterworts (Pinguicula)

California Pitcher Plant (Darlingtonia californica) Bladderworts (Utricularia)

ISBN 0 - 9 1 0 2 4 4 - 9 0 - 1