Carnivorous plant newsletter

CARNIVOROUS PLANT
NEWSLETTER

Journal of the International Carnivorous Plant Society

Volume 3 1 , No. 3 September 2002

CARNIVOROUS

PLANT

NEWSLETTEI

Journal of the Internationa
Carnivorous Plant Society
www.carnivorousplants.on

Volume 31, Number 3
September 2002

Front Cover: Heliamphora chimantensis x Heliamphora minor plants. Photograph by
Andreas Wistuba. Article on page 78.

Back Cover: Pinguicula moranensis ‘Libelulita’ flowers, photo by Barry A. Rice. Article
on page 83.

Carnivorous Plant Newsletter is dedicated to spreading knowledge and news related to carnivorous plants.
Reader contributions are essential for this mission to be successful. Do not hesitate to contact the editors with infor¬
mation about your plants, conservation projects, field trips, or noteworthy events. Contributors should review the
"Instructions to Authors" printed in the March issue of each year. Advertisers should contact the editors. Views
expressed in this publication are those of the authors, not the editorial staff.

All correspondence regarding dues, address changes and missing issues should be sent to the Membership
Coordinator at the ICPS. Do not send such correspondence to the editors. Checks for subscriptions and back issues
should be made to the ICPS in US funds. Dues for 2002 are $25.

ICPS, Inc.

PMB 330

3310 East Yorba Linda Blvd.
Fullerton, CA 92831-1709, USA
icps@camivorousplants.org

President

David Gray, email: david@camivorousplants.org
Carl Mazur, email: carl@camivorouspkints.org
Cindy Slezak, email: cindy@camivorousplants.org

John Brittnacher, email: john@camivorousplants.org, seedbank listed in this issue.

Vice President

Secretary
Seed Bank

Editors:

Barry A. Rice, PO Box 72741, Davis, CA 95617, USA, email: barry@camivorousplants.org
Jan Schlauer, Zwischenstr. 1 1, D-60594 Frankfurt, Germany, email: jan@camivorousplants.org
Page Layout: Steve Baker, email: steve@camivorousplants.org

Date of effective publication of the June 2(X)2 issue of Carnivorous Plant Newsletter: 6 June 2002.

The ICPS is the International Cultivar Registration Authority (ICRA) for cultivated carnivorous plants according to
The International Code For The Nomenclature of Cultivated Plants. Send relevant correspondence to the ICPS, Inc.

PUBLISHER: ICPS, Inc., Fullerton, California. Published quarterly with one volume annually. Desktop Publishing:
Steve Baker, 5612 Creek Point Drive, Hickoiy, NC 28601. Printer: Kandid Litho. Logo and masthead art: Paul
Milauskas. Dues: $25.00 annually. Reprints available by volume only © 2002 Carnivorous Plant Newsletter. All rights
reserved. ISSN #0190-9215.

65

Carnivorous Plant Newsletter

Contents

Abiotic Factors, Particularly C02 concentration, Affecting Carnivorous Plants from

the Eastern Shore of Maryland . - . 67

Book Review . 75

ICPS on the World Wide Web . 77

Heliamphora chimantensis, a New Species of Heliamphora (Sarraceniaceae)

from the ‘Macizo de Chimanta’ in the South of Venezuela . 78

New Cultivars . 83

Renewal Reminder! . 84

Seedbank . - . . . 84

Literature Reviews . - . . . 85

5th International Carnivorous Plant Conference: Call for Proposals . 86

An Interview with Dr. Rob Naczi About Sarracertia rosea . 87

Looking back: CPN 25 years ago . 94

Back issue sales: final notice . 94

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Contact: DAVID CRUMP Price list - S.A.S.E.

Volume 31 September 2002

66

Technical Refereed Contribution

Abiotic Factors, Particularly C02 concentration,
Affecting Carnivorous Plants from the Eastern
Shore of Maryland

Matthew McDermott and Douglas W. Darnowski • Department of Biology •
Washington College • 300 Washington Avenue • Chestertown, MD, 21620 USA •
Douglas.Darnowski@washcoll.edu

Keywords: Ecology: Maryland (USA), Utricularia, wetlands — Field studies:
Utricularia.

Submitted December 26, 2000

Abstract

Carnivorous plants, particularly bladderworts ( Utricularia spp.;
Lentibulariaceae), have long been known to grow on the Eastern Shore of
Maryland, but few if any studies have examined their growth in situ or in the lab¬
oratory along with the abiotic factors which affect their growth. From the scientific
literature, Great Swamp in Maryland (Kent County) clearly stands as an unusual
site for the Eastern Shore of Maryland, containing several species of bladderworts.
Upon investigation, four species of aquatic bladderworts were identified. Several
sites within Great Swamp, containing or lacking plants of Utricularia spp., were
routinely sampled to seek correlations between abiotic components of the local
aquatic environment, such as pH and [C02], and the presence of bladderworts.
Small samples of various Utricularia spp. were subjected to a variety of experi¬
mental treatments in the laboratory to examine further the factors identified in
Great Swamp. Moderate irradiance levels and elevated concentrations of dissolved
C02 both promoted the growth of U. intermedia and U. macrorhiza.

Introduction

Carnivorous plants have been an attraction for both botanists and horticultur¬
ists since before the time of Charles Darwin. These plants usually grow in nutrient-
poor environments, thriving in conditions which many species find daunting. The
occurrence of carnivorous plants in these environments relates to their special
traits, since to be considered carnivorous species must lure, trap, and digest prey to
obtain scarce nutrients especially nitrogen (D Amato, 1998). Factors besides the
availability of organic nitrogen may influence the growth of particular carnivorous
plants in particular environments. For example, just as irradiance levels may dic¬
tate the growth of understory plants versus canopy species, some biotic factors such
as the levels of inorganic nutrients or of pH might help to determine the presence
or absence of certain carnivorous plants.

Among carnivorous plants growing in the Eastern United States are the
American pitcher plants ( Sarracenia spp., Sarraceniaceae); the sundews ( Drosera
spp., Droseraceae); the Venus Flytrap ( Dionaea muscipula, Droseraceae); the but-
terworts ( Pinguicula spp., Lentibulariaceae); and the bladderworts (Utricularia

67

Carnivorous Plant Newsletter

spp.; Lentibulariaceae). Bladderworts can be found growing as terrestrial, sub-
aquatic, affixed aquatic, or submersed (suspended) aquatic plants (Taylor, 1989).

As indicated in the works ofTatnall (1946) and Sipple (1999), Utricularia spp.
seem to be the most common carnivorous plants found on the Eastern Shore of
Maryland, and, in particular, Sipple (1999) points to Great Swamp in Kent County,
Maryland, as a site containing an unusual abundance of bladderworts. This site
(which is actually a bog) consists of several interlinked channels of water created by
the mining of sphagnum peat. A portion of this area privately owned and under pro¬
tection by The Nature Conservancy, Echo Hill Camp and Echo Hill Outdoor School,
permitted access to part of Great Swamp.

Several sites were selected for routine measurements of water quality and for
observations of bladderworts (see Table 1). These sites varied with regard to the
amount of irradiance received, location near the bank or in mid-channel, and the
number of species which the sites contained. These sites were tested for a variety of
abiotic factors, both chemical factors such as phosphate levels and physical factors
such as the amount of ambient irradiance, to determine any relationship to the
presence or absence of bladderworts.

In previous studies from other geographic regions, communities of bladder¬
worts have been shown to exist under a wide range of temperatures and nutrient
conditions (Roberts, et al., 1985; Adamec & Lev, 2001). It has been shown by previ¬
ous research that nitrogen and phosphorous levels have a significant impact upon
the growth of bladderwort communities (Havens et al., 1999).

Non-destructive laboratory experiments were also conducted on small samples
of bladderworts from Great Swamp to further explore connections between levels of
illumination or dissolved carbon dioxide and the growth of bladderworts. Previously,
Adamec (1999) showed, and others confirmed (Camilleri, 1999) that elevated con¬
centrations of C02 in the aquatic environment accelerate the growth of Aldrouanda
vesiculosa (Droseraceae). This is another aquatic carnivorous plant, native to
Europe, Asia, Africa, and Australia, so levels of C02 in Great Swamp might influ¬
ence growth of bladderworts there.

Site # Description _ _ _

1 Mid-channel site next to fallen tree, bladderworts present among
Nymphaea odorata.

2 Bank site under heavy canopy. Largest community of bladderworts seen,
also large amounts of hornworts were observed.

3 Center of channel, no bladderworts present.

4 Bank site under heavy cover, receives afternoon sun. Bladderworts pre
sent.

5 Bank site with no cover, receives afternoon sun. Small numbers of
bladderworts present.

6 Bank site next to birdhouse, receives direct morning sunlight.
Bladderworts present among water lilies but not in nearby open water.

7 Bank site receiving evening sunlight, with heavy cover. No bladderworts

_ present. _

Table 1; Description of Sites in Great Swamp for Water Sampling

Methods

Using Taylor (1989), a monograph on the genus Utricularia currently accepted
as the standard taxonomic text, samples of Utricularia spp. from Great Swamp
were identified.

Volume 31 September 2002

68

Water samples were taken from several sites in Great Swamp, described in
Table 1, weekly for six weeks during June, July, and August of 2000. These samples
were tested as described in the LaMotte Monitors Handbook (Campbell, 1992).
Tests were performed for dissolved C02 (test kit #7297), toxic ammonia (#59100),

salinity (#7459), copper content (#10269), pH (#5090), nitrates (#3110), and phos¬
phate (#3114). New 50 ml polyethylene sample vials were used for each sample col¬
lection. At the pHs measured, all ammonia would have been in the form of NH4+,
and the data presented are for the appropriate test strip portion (Freshwater
Aqualab IV Mardel Glendale Heights IL).

For all experiments in the laboratory, clear plastic containers were used as
aquaria to contain bladderworts, and all replicates of a given experiment were con¬
ducted in identical containers. Plants were grown in a laboratory in which the tem¬
perature ranged from 20-25°C, approximately consistent with the range of aquatic
temperatures in which the bladderworts were found growing in Great Swamp.

To observe the effects of varying levels of irradiance, plants of U. intermedia, U.
macrorhiza, and U. gibba were grown individually in aquaria (three in parallel per
repetition; approximately 500 ml of doubly distilled H20 per aquarium to start) and
placed under constant irradiance provided by cool white-type fluorescent bulbs.
Controls received full strength irradiance (200-300 lux) while experimental plants
were grown with the same placement relative to overhead lights, but their aquaria
were covered by two layers of fiberglass window screen to decrease illumination
(60-80 lux). The experiment was run for three weeks per repetition for three repe¬
titions, and measurements were taken weekly of plant length, fresh weight, num¬
ber of traps produced, and the amount of irradiance received. The fresh weight of
the plants was measured after blotting excess water, starting with one plant per
aquarium or the same initial fresh weight if plants were small in the case of U. gem-
iniscapa in some experiments. Irradiance measurements were made using a Fisher
Scientific Dual Range Light Meter (Fisher Scientific, Pittsburgh, PA, USA).

Experiments were also conducted to determine the effect of varying levels of
C02 on the growth of bladderworts from Great Swamp. Controls were placed indi¬
vidually in separate containers and treated as for varied levels of illumination,
using the same species as in those experiments. Experimental plants were treated
identically to controls except that C02 generators assembled according to Camilleri

( 1999) were used to constantly bubble C02 through their aquaria. The experiment
was repeated three times for three weeks per repetition, and measurements were
taken weekly of plant length, the number of traps produced, fresh weight, and C02
levels.

Results

Three species of submersed aquatic bladderworts and one affixed species grew
in Great Swamp: U. gibba (submersed), U. geminiscapa (submersed), U. macrorhiza
(submersed); U. intermedia (affixed). Utricularia intermedia , U. macrorhiza, and U.
gibba had been found previously in Great Swamp by Sipple (1999). These plants
were identified mainly by leaf shape and bladder shape, and identification was con¬
firmed for U. macrorhiza and U. gibba by scape characteristics. These were the only
two species to flower during the period of observations.

Taylor (1989) clearly states that in spite of the similarities between the
European U. vulgaris and the American U. macrorhiza, they are to be referred to
separately, so his authority was followed. Utricularia gibba and U. macrorhiza
formed the great majority of plants found in the open swamp, where the seven sites

Carnivorous Plant Newsletter

69

for study were found. Utricularia intermedia was found in an area of much nar¬
rower and shallower channels along with those two species. Water quality was
examined less frequently in areas hosting U. intermedia, but in those sites the con¬
centration of carbon dioxide was similar to other sites containing a large number of
bladderworts, i.e. 14-20 ppm. Utricularia geminiscapa was only found occasionally,
in areas of shaded and open water. For laboratory experiments, U. macrorhiza, U.
gibba, and U. intermedia were used.

Water analysis at Echo Hill, Maryland, shows that the area of study had con¬
sistent levels for most of the abiotic factors examined at most sites studied during
the six weeks (June-August, 2000) of the study: pH (6.0; mean for all sites for 6
weekly measurements), nitrate (0.43-0.46 ppm), phosphate (0.25-0.29 ppm), salini¬
ty (1.02 ppt), copper (0.25-0.42 ppm), and temperature (24.4-24.7°C). Temperatures
were always within the range 23-26°C, even for the most extreme measurements
during this period. Levels of C02 and ammonia did vary among sites as shown in
Figure 1.

Stands of U. intermedia, U. gibba, and U. geminiscapa were found growing
under heavy to moderate cover along a boardwalk built through the swamp, where
the irradiance level varied from 2100-2200 lux from bright overcast to sunny con¬
ditions. Similar levels of illumination were found near the banks of channels. Both
U. macrorhiza and U. gibba were found growing in open water in the channels of the
swamp, though almost all stands of bladderworts observed were found with at least
partial shading by nearby aquatic non-carnivorous species. The irradiance level in
those areas and in areas with no floating cover, in which only a few plants of U.
macrorhiza and U. gibba could be found, varied from 20,000-21,000 lux.

All data from the growth experiments were normalized by dividing measure¬
ments for both control and experimental plants by the initial value for the control
plant in a given repetition. In Figures 2-3, data are shown without error bars
because the variation in the size of the plants which were available to start

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Site number

Figure 1: Levels of ammonia and dissolved carbon dioxide at seven sites in Great
Swamp. Mean±SD for six weekly measurements (not plotted) is 2.8 ppm for C02 and
0.73 ppm for ammonia. For descriptions of sites, see Table 1 .

Volume 31 September 2002

70

experiments was large. This introduces a large variation when experimental results
from separate repetitions are averaged, even after normalization. In spite of this,
the trends seen in Figures 2-3 for various experiments can also be seen when exam¬
ining data from individual experiments.

When the level of illumination was varied in
the laboratory, irradiance levels for the control
plants were around 200-300 lux and experimental
irradiance levels were at 60-80 lux. Controls of U.
intermedia and U. macrorhiza both showed weak¬
er growth in length with reduced irradiance lev¬
els. (Figure 2a, b). Utricularia intermedia also
showed a rather large increase in the amount of
traps produced at the control irradiance levels
(Figure 2c). U. gibba growth was unchanged by
changing irradiance levels (data not shown).

When the level of C02 was varied, levels for
experimental plants rose to around 13 ppm, com¬
pared to control levels at around 3 ppm. Elevating
levels of C02 enhanced growth of U. macrorhiza
and U. intermedia, as exemplified in Figure 3, dis¬
playing increases in trap number and fresh
weight for U. intermedia.

Discussion

The trends observed in the water quality
analysis fit what is expected for a site containing
bladderworts. Bladderworts can be found growing
in water within a pH range of 4-8 (Roberts et al.,
1985; Adamec & Lev, 2001), and in waters with
relatively low levels of nitrogen and phosphorus
(Havens et al., 1999), as were found at Great
Swamp where pH was typically 6 and nitrogen
and phosphorus were both less than 0.50 ppm.
Miniscule amounts of copper and ammonia were
observed, but as to whether or not they had an
effect on the growth of bladderworts is unknown
and is a subject for future testing. Carnivorous
plants most commonly grow in acidic environ¬
ments, and this was true for Great Swamp. The
levels of ammonia and phosphorus which were
measured were near or just below the recom¬
mended limits for the testing methods used, so
the principal point taken is that Great Swamp is
an area of moderate abundance for these nutri¬
ents.

Stands of bladderworts were most often
observed growing in moderately to heavily shaded
areas, most often crowded among Nymphaea
odorata (Nymphaceae), hornwort (Ceratophyllum
demersum, Ceratophyllaceae), and Nuphar adve-
na (Nymphaceae). Levels of irradiance in the
shade were not fully replicated in the laboratory

71

Figure 2: Effect of varied irradi¬
ance level (control plants approx¬
imately 200-300 lux; experimen¬
tal approximately 60-80 lux) on
the growth of bladderworts in the
laboratory. A. Change in length of
U. macrorhiza over three weeks.
B. Change in number of traps for
U. macrorhiza. C. Change in
number of traps for U. interme¬
dia. Mean of three repetitions.
Normalization and the lack of
errors bars are discussed in the
text.

Carnivorous Plant Newsletter

by control treatments, but plants grown in the laboratory at levels closest to those
preferred in nature showed increases in both plant length and trap production com¬
pared to plants treated with lower levels of irradiance (see Figure 2).

Bladderworts in Great Swamp probably prefer irradiance levels lower than
those of full sunlight, but very strong shade, represented by the experimental irra¬
diance treatment used in the laboratory, would be inhibitory of their growth. Even
those few plants which were found in full sun in open water grew among other non-
carnivorous aquatic plants which provided ample local shade. Production of irradi¬
ance intensities in the laboratory closer or equal to those of full sun should be exam¬
ined in future, using a method which offsets heat production to avoid confounding
experimental results. Higher levels of irradiance would also help to resolve any con¬
cerns about the levels used here having been at or below the compensation point for
photosynthesis. (Although it should be noted that the laboratory plants did grow
actively. )

Bladderworts growing both in the wild and in the laboratory at these irradiance
levels maintained a vibrant green color, while plants in the laboratory and in the
wild turned a reddish-yellow when subjected to higher irradiance levels. At Echo

Hill, U. macrorhiza was observed as the species
most tolerant to a range of irradiance levels, as
it could be found both in full shade crowded
among other plants or in full sunlight.
However, in full sunlight it was usually found
growing among other aquatics and was proba¬
bly not exposed to full sunlight. Utricularia
gibba was also observed in both areas, while U.
intermedia only occurred in shallow and shad¬
ed sites.

Utricularia gibba and U. macrorhiza in
full sun grew slightly submerged and did not
form partially-exposed surface mats. Such
mats did occur in the shade, however.
Inflorescences of both of these plants were
observed during the first three weeks of July
(U. macrorhiza), and during the last week in
July and the first week in August (U. gibba),
but only along bank sites that received partial
to full shade.

Levels of C02 in the wild were higher than
those achieved in the laboratory using a C02
generator. However, the strongest growth of
plants in the wild occurred at sites 1 and 2,
where the highest levels of C02, significantly
higher than other sites in the case of site 2,
were recorded (see Figure 1). However, levels in
the laboratory may have been depressed by
strong absorption of C02 by the rapidly grow¬
ing plants, or because the temperatures was
higher in the laboratory. It was observed that
U. intermedia, like Aldrovanda vesiculosa,
showed increased growth in length and trap
production due to increased C02 levels (see
Figure 3), probably due to the use of C02 in
photosynthesis (Adamec, 1999). The trends in

Figure 3: Effect of varied levels of
C02 on the growth of U. intermedia.
A: Change in number of traps. B:
Change in fresh weight. Control
level of C02 was approximately 3
ppm; experimental level was
approximately 13 ppm. Mean of
three repetitions. Normalization is
explained in the text.

Volume 31 September 2002

72

Figure 4: Utricularia habitat. Photo by Matthew McDermott.

growth, comparing control and experimental plants, were visible in the replicates
performed for each experiment, even though variations in the size of starting plants
made statistical analysis difficult.

Both U. macrorhiza and U. gibba seemed insensitive to C02 levels alone.
Perhaps both irradiance and C02 levels are important for the growth of bladder-
worts. U. gibba, the most widespread bladderwort in the world (D’Amato, 1998) and
perhaps the most adaptable, may not be affected strongly by the C02 levels used in
this study, but irradiance levels do seem important based on the placement of this
plant. Field observations by one author (DWD) of U gibba and U. muelleri, another
floating bladderwort, in the Northern Territory of Australia indicate a similar pref¬
erence there.

The distribution of traps and comparisons of trap production in response to
biotic versus abiotic factors were not attempted in this pilot study. However, such
work should be conducted in future in view of two papers: Knight & Frost (1991)
demonstrated that U. macrorhiza may change the numbers of bladders in response
to abiotic factors, while Friday (1989) showed that trap age and position are crucial
for evaluating the growth responses of bladderworts. Furthermore, Richardson
(2001) has recently suggested with a study of U. purpurea that bladders may also
have non-carnivorous importance in their functional ecology viz. a mutualistic rela¬
tionship with microinvertebrates surviving in the traps.

Utricularia macrorhiza and U. intermedia seem to be affected by both irradi¬
ance and the concentration of C02, and the different responses of these plants to
elevated concentrations of C02 might have explanations related to growth pat¬
terns — e.g. U. macrorhiza may grow to greater lengths at first to allow its modified
stems to photosynthesize more. Increased trap production may be more important
for the normally short U. intermedia, which seems to grow only in shade, than for
U. macrorhiza. Such factors, with the addition of biotic factors such as prey avail¬
ability, will be examined in future summers.

In future studies tests for potassium, calcium, and magnesium should be added
to the suite of tests used in this study as they have been shown to have an impact
upon the occurrence of U. intermedia and U. ochroleuca at sites studied in the Czech
Republic (Adamec & Lev, 2001). The experiments performed during the period con¬
sidered here did not allow for correlation between levels of some abiotic factors, such
73 Carnivorous Plant Newsletter

as ionic concentrations, and distributions of bladderwort species. This may be due
in part to the nature of the tests used, and further experiments to explore such vari¬
ables are planned for summer 2001. Measurement of photosynthetically active radi¬
ation using radiometric units will then be possible as well, due to recent acquisition
of equipment, and a broader range of C02 concentrations should also elucidate the
requirements of these species.

Acknowledgements

The authors gratefully acknowledge the financial support of the Thomas H. and
Barbara W. Gale Foundation (summer undergraduate research stipend to MM) and
of the Dean’s Office at Washington College (Faculty Enhancement Grant to DWD).
They also thank Echo Hill Camp and Echo Hill Outdoor School, and in particular
at those institutions, Andrew and Betsy McCown, for their generous permission to
study plants from Great Swamp.

Literature Cited

Adamec, L. 1999, The biology and cultivation of red Australian Aldrovanda vesicu¬
losa, Carniv. PI. Newslett. 28: 129-131.

Adamec, L. and Lev, J. 2001, Ecological differences between Utricularia ochroleuca
and U. intermedia habitats, Carniv. PI. Newslett., submitted.

Campbell, 1992, LaMotte Monitors Handbook, Lamotte Chemical Company,
Chestertown, MD, USA

Camilleri, T. 1999, An economical carbon dioxide generator, Carniv. PL Newslett. 28:
132-133.

Campbell, G. and Wildberger, S. 1992, The monitor’s handbook, LaMotte Chemical
Company, Chestertown, MD, USA.

D’Amato, P. 1998, The Savage Garden, Ten Speed Press, Berkeley, CA, USA.

Friday, L.E. 1989, Rapid turnover of traps in Utricularia vulgaris L., Ecology 80:
272-277.

Havens K.E., East, T.L., Rodusky, A.J., and Sharfstein, B. 1999, Littoral periphyton
responses to nitrogen and phosphorous: an experimental study in a subtropical
lake, Aquatic Botany 63: 267-290.

Knight, S.E. and Frost, T.M. 1991, Bladder control in Utricularia macrorhiza: lake-
specific variation in plant investment in carnivory, Ecology 72: 728-734.
Richards, J.H. 2001, Bladder function in Utricularia purpurea (Lentibulariaceae): is
carnivory important?, Am. J. Bot. 88: 170-176.

Roberts D.A., Singer, R., and Boylen, C.W. 1985, The submersed macrophyte com¬
munities of Adirondack lakes (New York, USA) of varying degrees of acidity,
Aquatic Botany 21: 219-235.

Schnell, D. 1976, Carnivorous plants of the United States and Canada, Lebanon
Valley Offset Company, Inc. Lebanon, PA, USA.

Sipple W.S. 1999, Days afield exploring wetlands in the Chesapeake Bay region,
Gateway Press, Inc. Baltimore, MD, USA.

Tatnall R.R. 1946, Flora of Delaware and the Eastern Shore, Intelligencer Printing
Company. Lancaster, PA, USA.

Taylor, P. 1989, The genus Utricularia - a taxonomic monograph, BPC Wheatons
Ltd., Exeter, Great Britain.

Volume 31 September 2002

74

Book Review

Schnell, Donald E. 2002. Carnivorous Plants of the United States
and Canada (2nd Edition). Timber Press; Portland, Oregon (tim-
berpress.com). 0-88192-540-3, 468 p. Hard cover, 16.5 x 23.5 cm
(6.5 x 9.25 in), $US39.95.

Reviewed by David O. Gray

All those interested in carnivorous plants have been eagerly
awaiting the revised edition of Dr. Donald Schnell’s monograph
on the carnivorous plants of North America. Happily, the 25 year wait was worth it;
the book is a masterpiece.

Its a hefty book, with a solid 373 pages of subject text, and over 200 good qual¬
ity photos throughout. The text is also illustrated by many clear botanical sketches
by David Kutt and other artists. For readers who are new to the carnivorous plant
world, the introduction includes a competent overview of the carnivorous plant syn¬
drome, relevant current scientific findings, and wetland ecology. Like the first edi¬
tion, there are chapters for each genera with an entry for each species, and while
the publicity says 45 species are discussed, I counted more that that. Each species
account has notes on cultivation, and with the author’s keen observations on habi¬
tats and many ravishing photos of plants in situ, hobbyists will find this book a
great aid in understanding their plants’ needs. The books concludes with a chapter
on possibly-carnivorous species, and one covering conservation issues, a glossary,
and a comprehensive bibliography.

Seemingly written for an American audience (although all measurements are
metric), readers from all parts of the globe will find this work valuable, with its
detailed discussions of habitat, conservation, and ecology, especially of the popular
Sarracenia. The text is not overly weighted with botanical jargon (there is a glos¬
sary of terms if you are rusty on your Latin), and the discussions are in a pleasing
vernacular. A variety of anecdotes add greatly to its readability and gives the book
charm. There are literature citations throughout, signaling this is a serious botany
as well.

Relevant nomenclature is sensible and logically discussed, and there will be
few surprises for most enthusiasts. Schnell is well know for his many papers on
Sarracenia (19 are listed in the bibliography), and here he clearly and logically dis¬
cusses the arguments on individual taxonomic controversies, and graciously refers
the reader to the works by authors with opposing views. The species and synonyms
are cross-referenced in the index and there is a list of the common names of
Sarracenia hybrids. The vexing Utricularia are clearly delineated for the amateur,
and similar species are cleverly paired in one description, which aids in learning
their differences.

One of the most valuable portions for this reader was the chapter on conserva¬
tion issues. Here is a sobering, if not depressing assessment from the author’s expe¬
rience of watching wetland habitats disappear over the last half of the century. He
details several conservation and recovery strategies, but the picture is not promis¬
ing. We can hope this will motivate stronger conservation efforts.

The book stumbles in its coverage of the western species. In discussing

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Carnivorous Plant Newsletter

Darlingtonia, various common inaccuracies are reprinted on that plant’s prefer¬
ences. The range map for this pitcher plant is wildly overdrawn and includes one fic¬
titious site and one well-known, but tiny introduced site. Other species’ western
ranges are misrepresented to various degrees, and the plants generally get short
shrift in print. These flaws are curious as the author cites the work of those such as
Rondeau who have great West Coast field experience. It is clear that Schnell’s
strength is in his long and great experience with the species of the American
Southeast. He could have profitably visited more sites in the Pacific states and
enlisted more input from colleagues in the West.

Interestingly, he states that Darlingtonia has no known enzyme production, but
includes its description with the known carnivorous species, rather than with the
“possible carnivorous species” such as certain species of Ibicella, Dipsacus, Catopsis,
and Capsella. Passi flora foetida is not mentioned at all, although it bears retentive
hairs, and homogenates from its tissues have been demonstrated to contain diges¬
tive enzymes (furthermore, it is native to Florida and the Southwest, and is a wide¬
ly introduced weed elsewhere).

Still, none of these shortcomings change the value of this important work. The
author has skillfully crafted a book that bridges important botanical work with
readable popular accounts. The cliche in this case fits: its should be on every book¬
shelf of those concerned with carnivorous plants. If you wish to inspire new genera¬
tions of botanists and enthusiasts, consider buying a second copy for donation to
your local botanical garden, nature center, or high school.

Propagators & Dealers of the highest quality carnivorous plants.

Sarracenia Pinguicula Nepenthes Drosera Dionaea

www.petflytrap.com

For ordering and product availability, please check our website or call toll free 1-888-437-1668
- Wholesale & Retail - Year Round Availability - Student Discounts - No Order Minimums - USA Only -

Contract tissue culture propagation available. Call for details.

Volume 31 September 2002 76

Writings from the Readership

ICPS on the World Wide Web

John Brittnacher • P.O. Box 72222 • Davis, CA 95617 • USA

Keywords: computers: internet.

If you have visited the ICPS web site recently you will have noticed the new
home page for your web site designed by iiberwebmaster and ICPS Vice President,
Carl Mazur. The main web site is produced and maintained by a team led by Carl.
The other members who work on the main site are Steve Venter, Pete Thiel, and
myself with the assistance of Ron Baalke, Steve LaWarre, Chris Teichreb, and Mike
Wilder. Barry Rice maintains the associated FAQ web site and Rick Walker and Jan
Schlauer keep up the Carnivorous Plant Database web site. Our design goal was to
have an uncluttered home page that is easy to use and would highlight what the
ICPS is all about. We did a series of “better one or better two” trials and discussed
the merits of each. It was fun watching the design change and improve over time.

Of course a new home page meant a new design for the 300-odd pages of the
web site. Even though I have done a large fraction of the site, I had forgotten how
much stuff is there. The process was slowed down because I had to re-read some of
the abstracts of talks and laugh again at the photos from the ICPS Conference 2000.
They are hidden in the New and Events section. And while I was checking that sec¬
tion I had to get caught up on what events were posted. The same thing happened
when I updated the archive of sample CPN articles. OK, so they are not as funny as
the conference banquet talk (what we could post on the web) but it had been a while
since I read Fernando Rivadavia’s travelogue of his expedition to find giant Genlisea
uncinata or about Chris Teichreb’s trek into Canada’s Northwest Territories. It also
reminded me that I had not gotten around to scanning in the article pictures we do
not have in electronic form. Maybe by the time this is in print they will be in elec¬
trons.

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Carnivorous Plant Newsletter

Technical Refereed Contribution

Heliamphora chimantensis, a New Species of
Heliamphora (Sarraceniaceae) from the ‘Macizo de
Chimanta’ in the South of Venezuela

Andreas Wistuba • Mudauer Ring 227 • 68259 Mannheim • Germany
Thomas Carow • Am Mustergarten 1 • 97702 Miinnerstadt • Germany
Peter Harbarth • Frankenweg 18 • 69221 Dossenheim • Germany

Keywords: new taxa: Heliamphora chimantensis, Venezuela.

Received: 8 February 2002

Introduction

During our January 2001 expedition to the tepuis of Venezuela (Wistuba et al.,
2001), we also explored parts of the ‘Macizo de Chimanta’, the Chimanta Massif in the
southwest part of the Gran Sabana. This huge massif covers a total area of 1470 km2
and is actually a cluster of tepuis including the central Chimanta Tepui itself. Their
peaks range in altitude from 1700 m (at the central part of the massif) to 2698 m (on
Eruoda Tepui). The ten tepuis that reach 2000 m above sea level cover an area of some
700 km2 in total (Huber, 1992). The size and the diverse altitudes of the Chimanta
Massif support numerous habitats including rivers, green valleys and forests as well as
rocky plateaus and moist savannas. Starting in the 1950s, various expeditions explored
the unique flora and fauna of this area. Many endemic plants and animals were dis¬
covered during these expeditions and certainly many more await discovery.

During this exploration we found a previously undescribed species of Heliamphora
most notable for its pitcher shape and the spoon shaped lids. This species seems to be
more closely related to Heliamphora tatei than to any of the other species described so
far from the Gran Sabana.

Heliamphora chimantensis Wistuba, Carow & Harbarth spec. nov.

Caudex ramosus; foliis caulescentibus vel rosulis; amphoriis in parte inferiore
infundibuliformibus, in parte media subventricosis, et in parte superiore cylindricis ad
leviter infundibuliformibus, longis 20-30 cm, latis (in parte superiore) 3.5 - 5 cm.

Inflorescentiis 3-5-floris, racemosis, ad 65 cm longis; flores nutantes; pedicellis 5-13
cm longis; petalis 4 lanceolatis, albidis vel pallide-roseis, 4-5.5 cm longis, 1.2-1. 7 cm latis;
staminibus ca. 20, 1-serialibus, filamentis 6 mm longis; antheris oblongo-lanceolatis, ca.
5 mm longis; ovario valde tomentoso; stylo glabro; stigmate 3 lobato; seminibus fuscis,
oblongis, ca. 3 mm longis, testa conspicue membranaceo-alata.

Rhizomes branching. Plants forming dense and often huge patches up to several
meters across.

Pitchers infundibulate in the lower half, slightly ventricose in the middle and cylin¬
drical to slightly infundibulate in the upper third (see Figure 1); pitchers 20 to 35 cm
long, 3.5-5 cm wide in the upper part; upper part of the pitchers completely glabrous on
the inner side; pitchers entirely green with deep red lids. Lid 1-2 cm wide and 2-2.5 cm
long, spoon-shaped, upright, ending with a sharp tip; the two lobes of the lid compressed
from the sides near the tip, often touching each other at the front, forming a quasi-hel¬
met; lobes are expanded in the lower part of the lid and narrowed sharply near the base;
the inner side of the lid with prominent irregularly shaped patches of glands, up to 5
mm across. Inflorescence 60 to 65 cm long, 3-5 flowers, peduncle slightly pubescent,
Volume 31 September 2002 78

pedicels 5-13 cm long. The lowest peduncle is the longest and bears a bract that fre¬
quently is transformed into a rudimentary pitcher. Tepals lanceolate, 4. 5-5. 5 cm long,
1.2-1. 7 cm wide, white to whitish-pink; ca. 20 stamens in 1 series, filaments 6 mm long,
anthers oblong lanceolate, 5 mm long, 1 mm wide; ovary 3 celled, pubescent, style
glabrous. Seed approximately 2 mm long, compressed, ovate, irregularly winged.

Specimens examined

Heliamphora chimantensis : Macizo de Chimanta, Section centro-oriental, 1921 m
a.s.l., N 05°16.672’; W 062°11.438’ Wistuba, Carow & Harbarth No. Chim 10.01.01/1,
holotype, flowering plant (VEN)

Heliamphora chimantensis : Macizo de Chimanta, Section centro-oriental, 1921 m
a.s.l., N 05°16.672’; W 062°11.438’ Wistuba, Carow & Harbarth No. Chim 10.01.0172,
isotype, flowering plant (VEN)

The two herbarium specimens nicely exhibited the characteristics typical of the
new species. The comments in this paper are based both upon these two specimens as
well as our field observations of many other plants in situ.

Figure 1 : Heliamphora chimantensis pitcher. Photograph by
Andreas Wistuba.

79

Carnivorous Plant Newsletter

Figure 2: Cluster of Heliamphora chimantensis pitchers. Photograph by Andreas Wistuba.

Figure 3: Heliamphora chimantensis pitcher appendage nectar patches (left inset) and
insect visitation; ant (left), wasp (right). Photographs by Thomas Carow.

Volume 31 September 2002

80

Distribution

Heliamphora chimantensis sometimes grows together with if. minor, the only other
species of Heliamphora recorded from Chimanta. As a consequence, hybrids frequently
could be found. However, H. chimantensis seems to prefer valleys growing at around
2000 m while H. minor also has been found in higher altitudes and usually prefers more
open habitats.

The H. minor plants on Chimanta are notable for the long and prominent bristles
inside the pitchers. This variant form of H. minor is actually fairly widespread and has
been found on many tepuis of the Chimanta Massif as well as on Aprada Tepui, but not
Auyan Tepui. Meanwhile, the typical form of H. minor is only known from Auyan Tepui.
The differences between these two forms of H. minor may merit further taxonomic
study.

Ecology

Heliamphora chimantensis plants were found growing exposed, in short vegetation
such as grasses, bromeliads, Xyris (Xyridaceae) or Stegolepis (Rapateaceae). These
plants do not grow taller than the H. chimantensis. In fact, we never found plants grow¬
ing in shaded locations. Often they grow near rivers in the valleys of Chimanta. In all
cases we found Heliamphora chimantensis growing together with Stegolepis ligulata,
and also often associated with Adenanthe bicarpellata (Ochnaceae) and various species
of Brocchinia (Bromeliaceae).

In comparison to the species found on the plateaus of the various tepuis, where the
surface usually is much more rocky and sandy and plants often can grow only on
‘islands’ of debris, highly limited in space, the moist savannah-like habitat H. chiman¬
tensis prefers allows the formation of huge dumps. We have visited many tepuis on this
and other expeditions, and had never before seen clumped Heliamphora colonies of com¬
parable in size to the ones typical of H. chimantensis. Vegetative reproduction seems to
play an important role as the seedling activity we observed was very low.

The lids of H. chimantensis bear strange, huge patches (up to 5 mm across) of
glands on their inner surface (Figure 3). They should be studied in more detail as they
seem to be involved in the attraction of prey. We observed ants and wasps being attract¬
ed by secretions of these glands but unfortunately our limited time on Chimanta did not
permit a detailed study. As we have observed with other species of Heliamphora, the
pitchers usually do not contain many captured insects.

Related species

The discovery of H. chimantensis on Chimanta and its characteristics came as quite
a surprise, as they clearly indicate that it is much more closely related to the southern
H. tatei, than to any of the northern species known to be growing in the Gran Sabana.
The flowers of all other species known from the Gran Sabana have 10-15 anthers, while
Heliamphora tatei var. tatei and H. tatei var. neblinae from the Amazon, and
Heliamphora chimantensis have about 20; however, while the anthers of H. tatei and H.
tatei var. neblinae are 7-9 mm long, those of H. chimantensis just reach 5 mm in length
(Maguire, 1978; Steyermark, 1984).

The nectar spoons of the other species from the Gran Sabana are shaped more or
less concave to form helmet-like structures under which nectar can accumulate, pro¬
tected from the frequent rainfalls. In H. tatei, as well as in H. chimantensis, the lids are
more upright. However, H. chimantensis protects its nectar production by the two lobes
of the lid which are bent forwards and are compressed from the sides to form a roof rem¬
iniscent of the helmets of the other north-eastern species. From the ones of H. tatei the
lids of H. chimantensis differ by the spoon-shape and the sharp contraction at the base.
In H. tatei the lids are only very slightly contracted at the base and rather rectangular
in shape.

81

Carnivorous Plant Newsletter

Hybrids

We observed numerous hybrids between Heliamphora chimantensis and the
Heliamphora minor on Chimanta. These hybrids can be easily distinguished from pure
Heliamphora chimantensis by the shorter pitchers, the red veins and the coarse bristles
inside the pitchers. Some of the hybrids’ pitchers showed deep red coloration, similar to
the form of H. minor growing on Chimanta. The helmet shaped lids of these show a
strong influence of Heliamphora minor. Apparently the hybrids are much more vigor¬
ous than either of the parent plants; we found patches of single clones measuring more
than 5 meters across (see Front Cover).

Etymology

This name was chosen as Heliamphora chimantensis because it is the only species
known to be restricted to Chimanta Tepui.

Discussion

The formation of huge clumps and the low seedling activity seem to be the conse¬
quence of the savanna-like habitat. In general, the seeds of Heliamphora clearly show
adaptations to dispersal by water. This works excellently on a “typical” tepui where lit¬
tle streams and shallow ponds carry the seeds to new small patches of debris, making
Heliamphora a kind of pioneer plant. However, the environment in the valleys on
Chimanta where we found Heliamphora chimantensis seems to be far more static, with
a well developed, continuous soil layer. The low vegetation might be too dense for
seedlings to germinate and grow. Those few seedlings which do survive to maturity can
develop into patches of huge size.

Although the large sizes of the clumps that Heliamphora chimantensis makes are
remarkable, they are not completely unprecedented. Heliamphora can multiply vegeta-
tively by division, but in most species the resulting clumps still remain small due to lim¬
itations of the detritus patches they grow in. Even so, we have observed similar, large-
clump growth patterns in Heliamphora hispida (Nerz & Wistuba, 2000), Heliamphora
heterodoxa (in the Gran Sabana) and Heliamphora nutans growing on the sandy foothill
area of Tramen Tepui.

As we already discussed in our publication on Heliamphora folliculata (Wistuba et
al., 2001) the pitcher-lids or so called nectar-spoons of Heliamphora are highly elaborate
structures for the attraction of prey. Apparently various strategies have been developed
by the different species of Heliamphora all using the lids in altered ways. Heliamphora
chimantensis with its gland-patches shows yet another way to use the lid-structure for
the attraction of insects.

Being morphologically fairly constant organs, the lids offer superb, yet previously
under-utilized, characters of taxonomic relevance (cf. Nerz &Wistuba, 2000; Wistuba et
al., 2001).

Literature

Huber, O. 1992, El macizo del Chimanta, Oscar Todtmann Editores.

Maguire, B. 1978, Sarraceniaceae ( Heliamphora ), in: The Botany of the Guyana
Highland Part-X, Memoirs of the New York Botanical Garden, 29: 36-61.

Nerz, J. and Wistuba, A. 2000, Heliamphora hispida (Sarraceniaceae), a New Species
from Cerro Neblina, Brazil-Venezuela. Camiv. PI. Newslett. 29: 37-41.

Steyermark, J.A. 1984, Venezuelan Guyana, Annals of the Missouri Botanical Garden,
71: 302-312.

Wistuba, A., Harbarth, P. and Carow, T. 2001, Heliamphora folliculata, a New Species of
Heliamphora (Sarraceniaceae) from the ‘Los Testigos’ Table Mountains in the
South of Venezuela. Camiv. PI. Newslett. 30:120-125.

Volume 31 September 2002

82

New Cultivars

Keywords: cultivar: Pinguicula moranensis ‘Libelulita’.

Pinguicula moranensis ‘Libelulita’

Submitted: 22 April 2002

In 1998, the Botanical Conservatory at the University of California, Davis, was
given an unsolicited gift of a crate of field collected plants, including many tens of
dormant rosettes of heterophyllous Pinguicula from southern Mexico. We acces¬
sioned these plants into the collection, and over the next several years observed
their characters. All the plants turned out to be Pinguicula moranensis, as we antic¬
ipated. The most striking specimen has a number of remarkable attributes, and is
being given the name Pinguicula moranensis ‘Libelulita’. This plant has already
been mentioned in the pages of this journal (Carniv. PI. Newslett. 29:2, p.55, 2000),
when one of us (BAR) reported on how the only specimen we had of this plant near¬
ly rotted to extinction.

In foliage, Pinguicula moranensis ‘Libelulita’ is not particularly different from
any of the other many clones of Pinguicula in cultivation, but in flower it is so strik¬
ing that some horticulturists have remarked that it nearly appears to be a new
species (see Back Cover). The five petals are large and square-tipped. They are over¬
all pink-purple, deeper so on the distal 2/3rds, and pale to near-white closer to the
petal bases. Where the petals fuse, they suddenly darken to a deep velvet red,
marked with a few white streaks on the lower-most petal. For us, the most amazing
feature of this cultivar is the bold petal venation. This venation reminds one of us
in particular (EMS) of the reticulated network of veins visible in the transparent
wings of dragonflies.

Since 2000, we have industriously propagated the plant vegetatively. (It is
amenable to leaf cuttings.) This plant will first be offered to the general public at
the annual University of California, Davis (USA) plant sale in October. We will sub¬
sequently distribute Pinguicula ‘Libelulita’ to specialists around the world.
(Specimens have already been sent to other horticulturists in the USA and Europe.)
Pinguicula moranensis ‘Libelulita’ should only be propagated by vegetative means
as there is no guarantee that seed progeny would maintain the cultivar’s subtle
characters.

The cultivar name was coined by Elizabeth M. Salvia on 18 October 2000, and
submitted by us for registration on 22 April, 2002. The Spanish cultivar epithet
means “little dragonfly”. Spanish was chosen to honor the country that houses
Pinguicula moranensis. According to ICNCP rules, either Pinguicula ‘Libelulita’ or
Pinguicula moranensis ‘Libelulita’ may be used for this cultivar. Additional pho¬
tographs of Pinguicula moranensis ‘Libelulita’ may be seen archived at
http://www.sarracenia.com.

— Barry A. Rice • P.O. Box 72741 » Davis, CA 95617 • USA *
barry@carnivorousplants.org; Elizabeth M. Salvia • 607 East 8th Street, #6A •
Davis, CA 95616 • USA • esalvia@rnother.com

83

Carnivorous Plant Newsletter

Renewal Reminder!

Do not forget to renew your membership promptly if your membership expires in
2002. Do determine what year your membership expires, look at the mailing label
on the envelope that contained your issue of Carnivorous Plant Newsletter. After
your name you will see 8 digits. The first four digits is your membership number.
The next four indicate the year your membership will expire. If it says 2002, your
membership will expire at the end of this year. (Unless you renewed very recently
and the number has not yet been updated.) If you need to renew, use the renewal
form included with this issue of Carnivorous Plant Newsletter to send your $25
membership fee to our Fullerton, California address. Note that you can use VISA or
MasterCard, and can renew for multiple years if you wish. Membership forms are
also on our web site (carnivorousplants.org) if you wish to use them.

International Carnivorous Plant Society Seed Bank

ICPS Seedbank* P.O. Box 72222 • Davis, CA 95617-6222 • USA

Darlingtonia californica — Oregon, USA
Dionaea muscipula
Drosera auriculata
D. binata
D. brevifolia
D. burmannii
D. capensis — narrow leaf
D. capensis ‘Albino’ — white flower
D. capensis — wide leaf
D. dielsiana

D. filiformis var. filiformis
D. gigantea
D. glanduligera
D. intermedia

D. intermedia — Florida, USA
D. intermedia — North Carolina, USA
D. intermedia — Rhode Island, USA

This is a partial list of the seeds available. A complete list is available online at the
ICPS web site, http7Avww.camivorousplants.org or by sending a self-addressed (stamped
if USA), envelope to the seed bank address.

Seed packets are US$1 each. Please include US$3 postage and handling for each order.
You may pay by cash, check, or money order in US$. Many members pay with cash. Please
make checks and money orders payable to “ICPS Seed Bank”.

The seed bank is a members-only benefit. The quantity of seed available to each mem¬
ber is 1 packet of each variety per month and 50 packets total in any 12 month period. Please
list alternative seed selections, as other orders will arrive before yours. If you have an e-mail
address, please include it so we can correspond should any issues arise. Seeds purchased
through the seed bank are intended for your personal use only and may not be sold.

The money raised by the seed bank is used by the ICPS to pay for seed bank expenses,
web site ISP charges, and ICPS educational and conservation programs. Donate seed and
get credit for free seed from the seed bank. Seeds of selected varieties are available free to
teachers for use in the classroom and to scientists and conservation organizations. It is ICPS
policy not to sell seed of plants protected by CITES Appendix I or the US Endangered
Species Act.

John Brittnacher, Manager • john@camivorousplants.org

D. macrantha subsp. macrantha
D. nidiformis
D. spatulata

D. stolonifera subsp. stolonifera
Nepenthes (maxima x fusca) x (spathulata x
spectabilis)

Pinguicula caerulea
P. lutea

P. primuliflora
Sarracenia alata
S. flava
S. leucophylla
S. psittacina
S. purpurea
Utricularia gibba
U. multifida
U. violaceae

Volume 31 September 2002

84

Literature Reviews

Araki, S. 2000, Isozyme Differentiation between Two Infraspecies Taxa of Utricularia
australis R. Br. (Lentibulariaceae) in Japan. Acta Phytotax. Geobot. 51: 31-36.

The author demonstrates that the plants identified as U. australis that grow in
Japan can be differentiated into two distinct groups (named f. australis and f.
tenuicaulis) on the basis of isozyme comparison. Different banding patterns have been
found for alcohol degydrogenase (ADH) and phosphoglucoisomerase (PGI), the
remaining 12 enzymes studied did not yield clear bands or different patterns. The
author suggests that the two Japanese groups investigated, corresponding to mor¬
phologically defined and geographically segregated taxa, may represent populations
resulting from at least two independent invasion events. (JS)

Casper, S.J. and Steiger, J. 2001, A new Pinguicula (Lentibulariaceae) from the pre-
alpine region of northern Italy (Friuli-Venezia Giulia): Pinguicula poldinii Steiger et
Casper spec, nov., Wulfenia 8: 27-37.

An issue of interest to some botanists is the apparent lack of Pinguicula lepto-
ceras/Pinguicula balcanica type butterworts in northeast Italy, especially since the
conducive climate, geography, and soils of the area suggest they could occur there.
Even though the region has been heavily and repeatedly botanized, this kind of
Pinguicula was never detected. As such, the recent discovery of a new species in this
region came as a great surprise. This paper reports on the detection and compares the
new plant to related species (i.e. P vulgaris, P. leptoceras, P. balcanica, P reichen-
bachiana, and P. fiorii).

The new species, Pinguicula poldinii , has relatively large flowers with the upper
petals reflexed. It is remarkable in having a somewhat irregular number of petals —
while most flowers have 5 petals, flowers with 6, 7, or 8 petals (and calyx lobes) are
apparently easily observed.

In some ways, this paper raises more mysteries than it answers. The plant’s
homophyllous nature sets it apart from the (heterophyllous) species most closely
related to it. It is also peculiar this plant has never been detected before, even though
in one area the plants were even growing on road banks! The paper proposes that
Pinguicula poldinii has recently colonized these road cuts from as yet undetected
nearby populations.

Some skeptical botanists may not believe that the new plant merit separate
species status, but the authors of this paper have a great deal of experience with
Pinguicula, and their opinions cannot be casually discounted. (BR)

Ellison, A.M. & Gotelli, N.J. 2001, Evolutionary Ecology of Carnivorous Plants. Trends
in Ecology and Evolution 16: 623-629.

The weakest point in this paper is the authors’ oversimplification of carnivorous
plant phylogeny. Contrary to statements made in the paper, Droseraceae and
Dioncophyllaceae (both Nepenthales) are not really independent lineages. They are
just not sister groups. It was only a single, eccentric researcher who has maintained
a monophyletic origin of all carnivorous plants until the 1980s. The sticky glands of
Droseraceae, Drorophyllaceae, and Dioncophyllaceae (as well as the remarkably sim¬
ilar glands in Nepenthaceae and the non-sticky genera in Droseraceae) are actually
homologous organs and they are not at all merely homoplasic (cf. CPN 26:34-38,
1997). Although both families probably belong to the same order Ericales (s.lat.), a sis-

85

Carnivorous Plant Newsletter

ter group relationship between Sarraceniaceae and Roridulaceae is by no means sup¬
ported unanimously by all gene sequence comparisons (only rbcL alignments suggest
such a close relationship). The proximity of Byblidaceae to Solanaceae was caused by
poor taxonomic sampling in the first (1993) genetic analysis. More recent data sug¬
gest a placement of Byblidaceae in Scrophulariales, not in Solanales. Cephalotaceae
are not placed in Geraniales but in Oxalidales, which is a separate lineage. It is not
sure if Roridulaceae are ancestral to Sarraceniaceae (it is not even clear if the two are
sister groups), and therefore it cannot be stated that adhesive traps are “simpler”
(from a phylogenetic perspective) than pitchers in Ericales.

Great emphasis is laid on a weak hypothesis that “most” carnivorous plants are
restricted to well-lit, nutrient-poor, waterlogged habitats. This generalization and the
largest part of the academic speculation derived from it are bound to collapse if but-
terworts (most of which dwell in shaded situations) or epiphytes (that do not inhabit
permanently waterlogged places) are considered. (JS)

Komiya, S., Toyama, M., Okita, S. & Shibata, C. 2001, Utricularia macrorhiza Le
Conte is Distributed in Northern Japan. Journal of Japanese Botany 76:120-122 (in
Japanese)

Unfortunately, no English abstract is provided in this paper but the distribution
map on p. 121 shows that most probably the plants treated as different Japanese
forms of U. australis in Araki’s paper discussed above are actually two different
species, viz. U. australis proper and the well-known north American and northeast
Asian U. macrorhiza. The latter species has not been recorded from Japan before,
probably because it had been confused with U. australis so far (even by respected
authorities such as Peter Taylor). Chorologically a southward range extension of U.
macrorhiza to northern Japan is plausible, and the present interpretation may be the
solution of a series of problems concerning these plants. In this light all records of
allegedly fertile specimens of “U. australis ” should be re-examined very carefully to
determine if they really belong to this species. ( JS)

5th International Carnivorous Plant Conference:
Call for Proposals

Dear fellow carnivorous plant enthusiasts,

Now that the 4th International Carnivorous Plant Conference at Tokyo has suc¬
cessfully concluded, the qualification process for the next conference is opened.

According to the geographic/chronological scheme for International Carnivorous
Plant Conferences, the next event should preferably be held in Europe, the Near
East, or Africa, in 2004.

All persons or societies interested in hosting the 2004 conference in or near the
intended region should contact me (Jan Schlauer, Zwischenstr. 11, 60594
Frankfurt/Main, Germany, <jan.schlauer@uni©tuebingen.de>) for further details.

Please note that suggestions for a venue alone (without a proposal to actually
organize the conference) cannot be considered in the qualification process.

Volume 31 September 2002

86

Writings from the Readership

An Interview with Dr. Rob Naczi
About Sarracenia rosea

Tim STEVENS • 4436 Blackwood Drive • Montgomery, AL 36109-3159* USA •
tstevensl954@mindspring.com

Keywords: Observations, taxonomy: Sarracenia purpurea , Sarracenia rosea — rec¬
ollections: Robert Naczi.

The following text is excerpted from an interview conducted by Tim Stevens on
2 October 2000. Stevens requested the interview from Robert Naczi in order to
learn more about the circumstances surrounding the description of Sarracenia
rosea as a new species (Naczi et al. 1999), as well as its conservation. Prior to the
publication of this paper by Naczi, Eric Soper, Frederick Case, and Roberta Case,
the plant was known as S. purpurea subsp. venosa var. burkii.

At the time of the interview, Naczi was an associate professor at Northern
Kentucky University. Since that time, he has moved to Delaware State University,
where he is curator of the Claude E. Phillips Herbarium.

Q: How did you become involved in studying Sarracenia rosea ?

RN: It’s actually an interesting story. It might not be exactly what most people
would suspect but I’m very interested in the whole community of arthropods that
live inside pitcher plants. There are some mites and insects that are actually able
to survive the plant. Most, as you know, get trapped and digested by pitcher plants
but there are some that live inside these pitchers and they live nowhere else. I’m
speaking of Sarracenia pitchers generally.

So I began a project as an undergraduate. I was looking at the mites that live
inside these pitchers and I was really fortunate to get a small grant and to make a
trip south in 1984. George Folkerts and some other folks from Auburn University
helped me a lot on that trip. So, they introduced me to these plants that I hadn’t
seen. I’d seen the northern purple pitcher plant but not this southern thing. Then,
when I entered graduate school, I did not pursue pitcher plants or their mites for
my doctoral research, but I did work on a group of plants that allowed me to do field
work in the southeastern United States. So I continued to collect from pitcher plants
and I did that in graduate school. I realized that the northern purple pitcher plant
looked quite a bit different to me than this plant on the Gulf Coast. So really, to
make a long story short, it was because of my work on the mites that live inside
these plants that I was taking a closer look at the plants themselves.

The more I worked on this the more I got interested in the plants themselves.
I had started out thinking that it had all been studied. But I realized, delving into
the literature, it hadn’t. So then I realized there was this potential to do something
here botanically.

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Carnivorous Plant Newsletter

Q: How did you become involved with Fred and Roberta Case?

RN: When I was a graduate student at the University of Michigan, I was fortu¬
nate to meet Fred and Roberta Case. They really took me under their wings and
were very generous with me, doing things like telling me about certain field loca¬
tions for various pitcher plant species. I’ve been in the field with them a few times.
So after I had started working on this project of just what is the status of these Gulf
Coast pitcher plants, I invited them to work with me. They had independently
noticed a lot of the differences I had, so we agreed that we’d work on this together.
That’s why they’re co-authors on the paper.

Q: They had been collecting and growing the species for some time in Saginaw?

RN: Yes, very much so. Their work in the greenhouse was a critical contribution
to realizing that Sarracenia rosea really is a species distinct from Sarracenia pur¬
purea because they had grown both Sarracenia purpurea and Sarracenia rosea
together in the same greenhouse, under the same conditions for years and years.
They didn’t need much convincing when I said, “Hey, I have evidence that these
things are different.” So when we put it all together we realized it was a compelling
case and that was another reason why I was glad to have them included.

Q: What role did Eric Soper play?

RN: He was an undergraduate when I was doing much of the study here at
Northern Kentucky University. He helped by measuring a lot of the specimens. It
was nice to have his contributions with all of his diligent work of measuring. He had
approached me about doing research and I described various projects to him in
which I was engaged, and he seemed to be interested in this one.

Q: When did you realize that this was probably a new species? How long did the
study actually last?

RN: Well, I didn’t realize — when I was an undergrad— that this plant was any
different. It was about 1987 or 1988 when I realized that the plant on the Gulf Coast
was different. I earnestly started work on it probably about 1990. In a way it’s
embarrassing that it took me so long but, on the other hand, it took me a while
because I wanted to do a thorough job. First, requesting all those specimens and get¬
ting them in from various herbaria, and visiting herbaria; doing the field work and
then all the measurements — it just took a long time.

The nice thing about doing all that field work is I was killing two birds with one
stone. I was working on the plant but I was also collecting mites and I’m continuing
to work both on the botany of Sarracenia and the mites.

The mites have been very little studied. People have overlooked them but it
ends up they’re a major component of this micro-ecosystem. But one of the reasons
I’m so interested in them, from the botanical standpoint, is that with many of the
Sarracenia species being rare I wonder what these arthropods are doing to or for
the plants.

There was one study published in the eighties by William Bradshaw from the
University of Oregon. He showed that the mosquitoes and the midge larvae that live
inside purple pitcher plants actually benefited the plants. When the mosquitoes and

Volume 31 September 2002

88

midges were present, the levels of nitrogen inside the pitchers were higher than
when the insect larvae were absent. So apparently, these insect larvae, by wriggling
through the prey remains, process them and release the nutrients faster.

Well, the mites are present in much higher numbers and they crawl through
the prey remains also and they fragment them. I don’t have any experimental evi¬
dence for this, but I hypothesize that the mites are actually beneficial to the plants
too. When I sample populations of the plants I find that the mites are almost always
present.

Q. What is your connection with George and Debbie Folkerts?

RN: I know Debbie and she’s done a lot of work on the moths. George has been
very, very generous with me in sharing his knowledge. So I really have high regard
for both of them. In fact, George’s paper, in 1982, in American Scientist, is one that
was really important to my undergraduate research. So I think it’s because of
George Folkerts, more than any other person, that I am pursuing these things.

Q: What are some features that distinguish Sarracenia purpurea from
Sarracenia rosea ?

RN: That’s easy. First, the thing to realize is that everything that has been
called Sarracenia purpurea from the Gulf Coast is this new species. It’s the only one
there in this group of Sarracenia. In other words, Sarracenia purpurea does not
make it that far south and west. So geography does it.

But that’s not very satisfying. If one is fortunate enough to be in the field dur¬
ing the blooming season, Sarracenia rosea has pink petals but Sarracenia purpurea
has maroon petals. The blooming season is quite short so most people would not be
there. So there are nice features that are present almost all year round. In terms of
the pitchers, the lip of the pitcher in Sarracenia rosea is much thicker than the lip
of Sarracenia purpurea. I give measurements in our paper but generally, just telling
people that it’s a thicker lip will do it.

Another thing that works very, very well — and this will work most of the year
because the plants are in fruit most of the year— is that the flowers and fruits of
Sarracenia rosea are quite a bit larger than Sarracenia purpurea and it becomes
especially conspicuous when you look at the relationship of flower size to height of
the scape. Sarracenia rosea has a large flower but a short flower stalk, or scape.
Sarracenia purpurea has a relatively small flower but a tall scape (see Figures 1,2).

So these are the most conspicuous differences. We found plenty of others.
Generally speaking, all aspects of the flower are larger, including petals. Petals are
longer and wider in Sarracenia rosea, and the pitchers tend to be larger, though
there’s a lot of overlap in that. So I really think I gave you the best differences, and
the ones that work best in the field as well as in the herbarium.

Q: Are these differences consistent in cultivation?

RN: Yes. That was important to us because we wondered, are some of these
things merely ecologic? So take it out of its geographic range and put it with
Sarracenia purpurea, and does it maintain those distinctions? Yes. It does for these
key differences.

89

Carnivorous Plant Newsletter

Q: How is Sarracenia rosea the most “genetically divergent”?

RN: It wasn’t the most divergent member of the genus. Mary Jo Godt and Jim
Hamrick from the University of Georgia were looking at genetic diversity within the
Sarracenia purpurea complex. So they had quite a narrow scope, but within that
group, the one plant that stood out the most — it was the most different genetically
from all the others and that’s what we mean by genetically divergent — was
Sarracenia rosea. The genetic difference between it and the next most genetically
closely related member of the Sarracenia purpurea complex, was as much as or
greater than a lot of investigators have found for separate species. Basically, it was
the most different among any of those that they looked at, and they looked at four
taxa — Sarracenia rosea, the northern Sarracenia purpurea subsp. purpurea, the
mid-coastal Sarracenia purpurea subsp. venosa, and the mountain Sarracenia pur¬
purea subsp. venosa var. montana.

Q: What about the ranges of Sarracenia rosea and Sarracenia purpureal

RN: The thing that worries me is the map that I provide. If one just looks at that
they may get a false sense that this plant is more common than it is. For instance,
we found herbarium specimens from two populations in Georgia, but those are
unknown presently. The plant is most likely extirpated from Georgia and a lot of the
mapped locations are gone. So even though the region from the mid-Florida pan¬
handle west to Mississippi is kind of thick with dots on our map, a lot of those dots
are no longer there — a lot of those populations are no longer there. I’m very con¬
cerned about the conservation of the plant.

Q: In Alabama, is the plant mostly found in Baldwin and Mobile counties?

RN: Yes. Baldwin and Mobile are — in terms of Alabama — the only places where
this plant is fairly frequent. It’s still a rare plant. So I think we need to be worried
about it and I mention in the paper two instances that I myself witnessed of poach¬
ing of the plants.

Q: Are the plants well distributed throughout the known sites or do two or three
sites have most of the known plants?

RN: That’s just what I was going to say, that a lot of the sites I found when I
was doing all that field work for all those years and really scouring the areas, I
would find five, six, seven, eight, nine, ten, a dozen. Very few sites have what I would
call large populations. So I’m hoping the plant is a lot more common than we real¬
ize. I think there needs to be a status survey done. But in my experience, even in
areas where I know it is and it’s good habitat, I don’t find much of it.

Q: Are the good sites on public or private lands?

RN: Both. Like two of the best populations I know — one of which is the type
locality in the Apalachicola National Forest in Florida. Then, in Alabama, the places
I’ve seen the plant are all private land.

Volume 31 September 2002

90

Figure 1 : Pressed specimens of Sarracenia purpurea (left) and Sarracenia rosea (right).
Photograph compliments of Robert Naczi.

Figure 2: Sarracenia rosea in Florida. Photograph by Barry A. Rice.

91

Carnivorous Plant Newsletter

Q: Are these burned regularly?

RN: One of the places definitely is.

Q: What is the typical Sarracenia rosea habitat like? Is it always found with
Sarracenia leucophylla ?

RN: That’s the real indicator. I’m not saying that every place you find
Sarracenia leucophylla, you find Sarracenia rosea. In fact, no. But if Sarracenia leu¬
cophylla is there one could get out and really look around.

Q: What are the greatest threats?

RN: I really think it’s habitat destruction. It’s not the over collecting or poach¬
ing. I mean those are threats and I’d rank them as serious threats, but the most
serious threat, in my experience, is this destruction of habitat because of the rapid
development. I myself have seen pitcher plant habitats disappear in the relatively
few years that I’ve been at it. The boom of development, especially right along the
coast, is just astounding. So I see that as the worst. Fire suppression, I would say,
would be the second most serious threat. Then I would rank poaching as the third
most serious threat. Again, we need a status survey but, in my experience, those are
the most serious threats prioritized.

Q: How do other workers feel about recognizing Sarracenia rosea as a distinct
species?

RN: I know there will always be differences of opinion. There will be a lot of peo¬
ple who’ll say I’m just a splitter or I just wanted to describe a new species. I think
the best way to go is to document diversity and I think if we don’t recognize some¬
thing that’s truly distinct as a species, we do an injustice to it because, especially for
conservation purposes, these things don’t get as much priority when they’re vari¬
eties or subspecies as when they’re species. In my opinion we provide many reasons
why this is distinct as a species: the morphology, we cite the genetic work of Godt
and Hamrick, we have the greenhouse common growth experiments that the Cases
did. I know people will disagree with me. The best I can do is lay my cards on the
table, show the evidence of it, and let people make their decisions.

Papers mentioned in this interview:

Bradshaw, W.E. and R.A. Creelman. 1984. Mutualism between the carnivorous

pitcher plant and its inhabitants. Am. Midi. Nat. 112: 294-304.

Folkerts, G.W. 1982, American Scientist, The Gulf Coast pitcher plant bogs, 70: 260-

267.

Godt, M.J.W & Hamrick, J.L. 1999, Genetic Divergence Among Infraspecific Taxa of

Sarracenia purpurea, Sast. Bot., 23: 427-438.

Naczi, R.F.C., Soper, E.M., Case, F.W., & Case, R.B. 1999, Sarracenia rosea

(Sarraceniaceae), A New Species of Pitcher Plant From The Southeastern

United States, Sida, 18(4): 1183-1206.

Volume 31 September 2002

92

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93

Carnivorous Plant Newsletter

Index of Nomenclatural Novelties in This Issue

Heliamphora chimantensis . -78

Looking Back: CPN 25 years ago

Steve Rose supplied an excellent five page review of tuberous Drosera of
Western Australia — fine reading for those who cultivate plants in the group.
Meanwhile, Don Schnell found a street named “Sarracenia”, appropriately enough
in Sarracenia alata territory in Escatawpa, Mississippi. The discovery drove him to
poetic musings: “...perhaps it would be appropriate one day if on this road someone
built a home for aged and retired CP botanists who in senior and calmer years,
somewhat like old warriors looking back on ancient battles, might sit on a wide
veranda and discuss old times in fields long grown over....”

Back Issue Sales: Final Notices!

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After 2002, we will only stock current issues of the journal!

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CPN Back Issue Clearance Sale

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Volume 31 September 2002

94