CARNIVOROUS PLANT NEWSLETTE1
Journal of the International Carnivorous Plant Society
Volume 35, No. 2 June 2006
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CARNIVOROUS
PLANT
NEWSLETTER
Journal of the international
Carnivorous Plant Society
www.carnivorousplants.org
Volume 35, Number 2
June 2006
Front Cover: Darlingtonia californica and Pinguicula macroceras subsp. nortensis in
California. Read about Darlingtonia in Canada on page 36. Photograph by B. Rice.
Back Cover: The white-petalled taxon of Drosera microphylla in the wild. Note the
vibrant red colouration of the plants. The plants are growing in moss-covered skele¬
tal soil over granite, with Borya sp. and D. glanduligera. Note the white petals of the
D. microphylla flowers are visible in bud, but no flowers were open that day due to
cool and cloudy conditions. Photograph by Robert Gibson. Article on page 39.
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 shouldJrej^gaLtQ the Mernhpr'*hip-
Coordinator at the ICPS. Do not send such correspondence to thejeditors. Cjhm^(^r|Sptjsfriptidu^ciitd^£k issues
should be made to the ICPS in US funds. Dues for 2006 are $25. j i I D R A R v
President
Vice President
Treasurer
Secretary, Seed Bank
Boar d Member
Board Member
JUN 2 2 2006
ICPS, Inc.
PMB 322
1 564-A Fitzgerald Drive
Pinole, CA 94564-2229, USA
icps@camivorousplants.org
NEVV Twr\o
Carl Mazur, email: carl@camivoroipplants.ogQT/\N !C AL G ARD EN
Cindy Slezak, email: cindy@carniviE>rouspIanto.org-^ —
Richard Myers, email: richard@camivorousplants.org
John Brittnacher, email: john@camivorousplants.org, seedbank listed in this issue.
Chris Teichreb, email: chris@camivorousplants.org
Steve LaWarre, email: stevel@camivorousplants.org
Editors:
Barry A. Rice, P.O. 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 March 2006 issue of Carnivorous Plant Newsletter: 22 March 2006.
The ICPS is the International Cultivar Registration Authority (1CRA) 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., Pinole, California. Published quarterly with one volume annually. Desktop Publishing:
Steve Baker, 5612 Creek Point Drive, Hickory, NC 28601. Printer: Kandid Litho. Logo and masthead art: Paul
Milauskas. Dues: $25.00 annually. © 2006 Carnivorous Plant Newsletter. All rights reserved. ISSN #0190-9215.
34
Carnivorous Plant Newsletter
Contents
Seedbank -
Darlingtonia californica (Sarraceniaceae) and Sarracenia purpurea (Sarraceniaceae) —
profiles of a refugee and a recluse in British Columbia -
Election announcement -
White-petalled Drosera microphylla Endl. from near Esperance, Western Australia -
Heliamphora exappendiculata , a clearly distinct species with unique characteristics - -
Naphthoquinones content of some sundews ( Drosera L) -
Plumbagin content in Aldrovanda vesiculosa shoots -
Looking back: CPN 25 years ago - : —
Book review -
Literature reviews -
Unexpected hybrids of spectacular bladderworts species -
■35
36"
■38
-49
■52
55
56
•57
59
International Carnivorous Plant Society Seed bank
ICPS Seedbank • P.O. Box 71 • Ashland, OR 97520-0003 • USA
Drosera auriculata
D. binata
D. binata — Coromandel, NZ
D. burmannii
D. capensis ‘Albino’ — white tlower
D. capensis— narrow leaf
D. dielsiana
D. fdiformis var. filifonnis
D. glanduligera
D. intermedia — Cuba
D. intermedia — New Jersey, USA
D. intermedia — North Carolina, USA
D. intermedia — Rhode Island, USA
D. intermedia — Venezuela
D. peltata
D. rotundifolia — Alaska, USA
D. rotundifolia — Washington. USA
/). spatidata
D. stolonifera subsp. porrecta
D. tokaiensis
Proboscidea louisianica (noncarnivorous)
Sarracenia flava
S. leucophylla
S. oreophila — ESA/CITES, USA sales only
S. purpurea subsp. purpurea
S. purpurea subsp. venosa
S. rubra — infraspecific hybrid
S. rubra subsp. jonesii — ESA/CITES. USA
sales only
Note: The seedbank address has changed! The new address is above!
This is a partial list of the seeds available. A complete list can be obtained online at the ICPS
web site, http://www.camivorousplants.org/ or by sending a self-addressed, stamped (if USA), enve¬
lope 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 member is 1
packet of each variety per month and 40 packets total in any 1 2 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.
You are encouraged to stock the seed bank with seed produced by cultivated plants. The ICPS
policy on wild seed collection is on line at the ICPS web site. Cultivation-produced seeds of species
protected by the US Endangered Species Act are distributed within the US only, and in accordance
with the ICPS's US Fish &Wildlife Service permit.
Donate seed and get credit for free seed from the seed bank. Seeds of selected varieties are avail¬
able free to teachers for use in the classroom and to scientists and conservation organizations. It is ICPS
policy not to sell internationally seed of plants protected by CITES Appendix I or the US Endangered
Species Act.
John Brittnacher, Manager •john@carnivorousplants.org
Volume 35 June 2006
35
Writings from the Readership
Darlingtonia californica (Sarraceniaceae) and
Sarracenia purpurea (Sarraceniaceae) — profiles of a
REFUGEE AND A RECLUSE IN BRITISH COLUMBIA
Chris Teichreb • 39 Dynes Street * Red Deer • AB T4R 3C3 • Canada • cteichreb@hotmail.com
Keywords: observations, non-native species: Darlingtonia californica , Sarracenia purpurea ,
British Columbia (Canada).
Darlingtonia californica (Torrey) and Sarracenia purpurea subsp. purpurea (L.) are both
hardy pitcher plants commonly found in carnivorous plant enthusiasts’ collections. Darlingtonia
is restricted to suitable habitats in a range extending from Oregon to northern California, but it
is capable of surviving along the southwestern coast of British Columbia (B.C.), where the cli¬
mate is virtually identical to its native range. Sarracenia purpurea subsp. purpurea is cold toler¬
ant and could grow in any suitable location within B.C. However, only Sarracenia purpurea
subsp. purpurea is naturally found in B.C., and even then only within a restricted area. Further
discussions of both genera, as they occur in B.C., are presented below.
So how did this all begin, or rather, where did I come in? Adolf Ceska, a Victoria (B.C.)
botanist and editor of Botanical Electronic News (BEN) posted a question to the carnivorous
plant listserv e-mail discussion group asking if anyone knew whether the Darlingtonia found in
Pacific Rim National Park, Vancouver Island. B.C. was a natural, disjunct population, or whether
it was introduced. I casually replied that I knew it was introduced, as I knew the person who
introduced the seed to this bog. The next thing I knew, Adolf was asking me for an article for
BEN and Barry Rice was asking me for an article for Carnivorous Plant Newsletter!
The saga began in 1999. when a carnivorous plant enthusiast1 from Vancouver went on
vacation to Oregon and collected a few ripe Darlingtonia seed pods. On a subsequent visit to the
Pacific Rim National Park's Shorepine Bog2, he randomly distributed the seeds as he walked
along the boardwalk. The enthusiast told me about the location, but attempts I made at locating
the plants in early-summer 2001 were unsuccessful, likely due to the small size of the plants, and
a spouse who was more interested in the ocean than a mosquito-infested bog (can you imagine?).
So, it was with a bit of surprise that I read Adolf’s question regarding the Darlingtonia. In
subsequent conversations with other botanists and naturalists. Matt Fairbams provided me with
photos of the plants he had taken in 2004 (see Figures 1 and 2; note the presence of Drosera
rotundifolia for size reference). The small size of the plants indicated that these were indeed the
result of the seed spread in 1999. Matt Fairbams indicated that the leaves were small compared
to natural populations in Oregon, and that the plants had not flowered in 2003. This is not sur¬
prising, given that Darlingtonia can take more than six years to reach flowering size, and even
after that stage plants continue to increase in pitcher size for many years. Although some think
that this population may eventually fail, I believe that it is well-established, and if left undis¬
turbed. will persist into the future. Currently, there are four known patches of Darlingtonia. but
others may easily be hidden among the native vegetation in the bog.
While some people may think that the introduced population of Darlingtonia is a natural
occurrence in B.C.. in fact the only native carnivorous pitcher plant is the common Sarracenia
purpurea subsp. purpurea. This species is widespread across much of Canada and is even the
provincial Bower of Newfoundland. The Rocky Mountains act as an effective barrier to the
'Who wishes to remain anonymous.
2At the request of Park officials, I have not revealed the exact location of the plants due to fears
that people may leave the boardwalk to search for these plants.
36 Carnivorous Plant Newsletter
Figure 1: Juvenile Darlingtonia californica in British Columbia. Photograph by Matt
Fairbarns.
Figure 2: Tiny pitchers of Darlingtonia californica in British Columbia. Photograph by
Matt Fairbarns.
Volume 35 June 2006
37
pitcher plant's spread westward, but a large portion of northeastern B.C. is east of the Rockies.
It is here that Sarracenia purpurea subsp. purpurea has been detected.
A single location for the genus in B.C. was reported by Krajina ( 1968), and was given as
being near Jackfish Creek, south of Fort Nelson. (I repeat this locational data here for two rea¬
sons; first, the location was reported in Krajina’s paper and second, the location is difficult to
access and is now an ecological reserve). Numerous plants were reported as occurring in the
location. Lamb (1989) noted a second location existed near the Fort Nelson location. Recent
searches of the University of British Columbia (UBC) herbarium records and of the scientific lit¬
erature have not revealed any new locations. Thus, it appears that these two populations are dis¬
junct from the rest of the range of Sarracenia purpurea.
A single plant from the Krajina population was collected sometime in the past (perhaps in
the early 1980s) and introduced into cultivation. At one point I was fortunate enough to obtain a
division of this plant, but it was not as robust as some of my other Sarracenia purpurea subsp.
purpurea. The one time it produced a flower stalk, the flower was consumed by a greedy slug
before opening, though it appeared to be the typical dark red flower.
These plants still pose many unanswered questions. Why does S. purpurea subsp. purpurea
apparently only occur in a very restricted area even though there is suitable habitat in many other
locations in northeastern B.C.? Why is Darlingtonia naturally restricted to Oregon and northern
California, when the climates in the southern coastal region of B.C. and the west coast of
Washington are so similar and can support populations of Darlingtonia ? (There are even the
occasional serpentine seeps with Pinguicula in B.C., two excellent indicators for Darlingtonia
in California!) Things to mull over in a mosquito infested bog.
References:
Krajina, V.J. 1968. Sarraceniaceae, a new family for British Columbia. Syesis, 1; 121-124.
Lamb, R. 1989. Sarracenia purpurea in Western Canada. Carniv. PI. Newslett. 18. 7-8.
Board Member Elections
The end of this year will mark the end of terms of office for four ICPS board members:
Carl Mazur (President), Chris Teichreb, Steve LaWarre. and Richard Myers. The other board
members, as well as all the additional volunteers responsible for operating the ICPS, extend their
thanks to these officers for the work they have done during their terms of office.
The four officers facing the ends of their terms have not yet indicated if they are interested
in running for another term. In any event, all members of the ICPS are encouraged to run for a
board position in the ICPS. If you would like to participate in the workings of the ICPS, please
email a 250 word election statement to Barry Rice (contact information is on the inside cover of
Carnivorous Plant Newsletter). Your election statements must be received by 1 August. 2006. In
your statement, you may wish to note your experience, why you want to run for office, and what
you wish to accomplish. Your statements will be printed in the September issue.
The ICPS is a growing, active organization. In order to participate as a board member,
email access is essential. Furthermore, board members are expected to work hard! So while we
welcome and encourage your interest as a board member, remember that it is more than a van¬
ity post! Consider running for office — your energy is appreciated and needed!
38
Carnivorous Plant Newsletter
International Correspondent
White-petalled Drosera microphylla Endl. from near
Esperance, Western Australia
Robert Gibson • 12 Roath St • Cardiff • NSW 2285 • Australia • rgibson@dipnr.nsw.gov.au
Keywords: ecology — observations: Drosera microphylla.
In the coastal granite hills near Esperance (33'49'S, 121'52’E), Western Australia grows a
variant of Drosera microphylla Endl. that has white petalled flowers and orbicular leaves, and
thus differs from the typical form of the species. In this paper this variant is described and com¬
pared with published descriptions.
The coast near Esperance is dominated by three different types of environment: coastal sand
dunes composed of either quartz sand or carbonate sand of biogenic origin; a subdued low
plateau with laterite weathering products; and low, often rounded granite hills that rise above the
landscape. Each type of landscape supports a suite of carnivorous plant species, but for now we
focus on the granite hills. Whilst the hills are modest by any standard, they occur at or very close
to the coast and form enough of a physical barrier to deflect on-shore winds and generate more
cloud and rain than other parts of the coast, even if only slightly so. Water runs quickly down the
bare granite slopes and is gradually channelled into the headwaters of numerous small intermit¬
tent streams. On the edge of bare granite slopes soil formation is minimal. The thin soil is often
covered by moss, and a selection of drought-deciduous forbs and annual herbs. It is in this envi¬
ronment that this taxon of D. microphylla occurs, and often does so in great abundance (see Back
Cover).
The soil dries out quickly in late spring, and the tubers1 of this sundew are nestled snugly
at least 10 cm deep in the soil, where possible, or nestled into any available clefts in the granite
surface where the soil is thinner. In autumn the weather cools and the incidence of rainfall
increases. Growth commences from the tubers and by mid to late autumn growth emerges at the
soil surface. Juvenile plants form a rosette of spathulate leaves but larger plants bypass this stage
and immediately begin to form an erect stem with orbicular cauline leaves. A conspicuous char¬
acteristic of the taxon is the vibrant red colouration of the leaves and stem, which is made all the
more eye-catching by the density at which these plants grow together.
The plants grow throughout the winter and into mid-spring when a surprisingly small num¬
ber of plants begin to form a terminal inflorescence. I have observed plants of this taxon flower¬
ing in October and November in the wild. The sepals are large, smooth and have a metallic lus¬
tre, which is typical of this species. The petals are white with an often variable amount of red
pigment on the outside base of the petals; at the flower’s maturity, this colouration results in a
bulls' eye feature that may assist in attracting pollinators (Figure 1 ).
In late spring to early summer the soil begins to dry out, and the plants senesce and become
dormant. The timing of dormancy is variable, and appears to at least partially correlate with soil
depth — plants in thin soil senesce before those in deeper or more sheltered soil. This can be seen
where plants on the edge of a mossy soil area are dormant whereas those in the deeper soil near
the middle are still alive.
'As the storage and resting organs in subgenus Ergaleium are covered by a leaf-derived, readily
detachable envelope, they must not be called tubers (in which the adnate skin is not derived from
leaves and detachable only by force or after cooking) but are instead what the botanist techni¬
cally calls corms, cf. the analogous situation in Crocus or Gladiolus. — ed. (JS)
Volume 35 June 2006
39
Figure 1 . Detail of an open flower. The red pigment on the outside base of the petal is
transmitted through to the inner centre, giving it a dark pink tinge.
Following the format in Marchant and George (1982) this taxon is described as:
Herb with a tuber. Stem erect, 5 to 18 cm tall, glabrous, weakly llexuous, with several
bract-like prophylls, 3 to 6 mm long at the base. Leaves alternate, rarely with secondary
leaves in the upper axils; lamina orbicular, deeply cupped, 3 mm diameter, peltate, on
petiole 8 to 20 mm long. Inflorescence a small panicle, 1 to 5-flowered; pedicels 5 to
30 mm long. Sepals 5, narrowly obovate, apex obtuse, deeply concave 7 to 10 mm long,
denticulate, glabrous, iridescent, olive green. Petals 5. obovate. 8 mm long, 7 mm wide,
unscented (to the human nose), white with the abaxial surface of the petal base often
deep red. Ovary, 1, three-locular, red, ovoid, glabrous, 1.2 mm diameter and 0.8 mm
long. Styles 3, 2 mm long, flattened, multiply divided into many terete segments form¬
ing a low dome over the ovary. The basal third of the styles are red, with the remainder,
including the stigmas, white. Stamens, 5, erect, 4 mm long. Filaments 3 mm long by 0.6
mm wide, pale pink, flattened in cross-section. Anthers, 5, 0.8 mm long, pollen yellow.
Bracteoles spathulate, 1 to 2 mm long, olive green to red, glabrous, with entire margins:
the apex is often cupped. Seeds oblong, dark grey, flattened, reticulate surface, to 1.8
mm (including membranous apical ends) long by 0.3 mm wide by 0.2 mm thick, the
ends extending into often-tlexuous wings to 0.6 mm long (Figure 2).
Drosera microphylla is known to be a polymorphic species (Diels, 1906: 121; Lowrie, 1987:
64). This species is endemic to the south west of Western Australia, where it grows in laterite soils
in Jarrah forests. Over the majority of its range it consists of olive-green plants with sub-orbicular
leaves, and red or orange-coloured petals. A summary of key features for different members of the
complex is presented in Table 1 (below):
In this article I have provided a description of a distinctive member of the attractive D. micro¬
phylla complex. This taxon appears to possess characters that are unique in the complex: an over¬
all red colouration to the plants, deeply cupped cauline leaves and white petals. At this stage the
taxonomic status of this entity is not yet known, but the plants appear to form a coherent popula¬
tion, with consistent characters and grows apart from the other members of this complex. Further
taxonomic study into this complex appears warranted, and would likely be most rewarding.
40 Carnivorous Plant Newsletter
Figure 2. Botanical illustration, by R. Gibson, of the white-petalled taxon of Drosera
microphylla. A: Whole plant in flower; B: prophyll; C: cauline leaf; D: bracteole; E: sepal;
F: petal, with the red base indicated; G: stamen — adaxial view; H: stamen — abaxial
view; I: open flower; J: gynoecium; K: mature seed. Scale bar in all cases is 1 mm.
Volume 35 June 2006
41
White-
petalled plant
Diels 1 906 var.
microphylla 1
Diels 1906 var.
macropetala 1
Lowrie
1987
Orange-
petalled plant
Cauline leaf shape
& diameter x depth
Orbicular
3 x 1.5-2
Suborbicular
1.5-3 x 1.5-3
Suborbicular
1.5-3 x 1.5-3
Re ni form
3-3.5 x 1
Orbicular
2 x 1
Cauline leaf
curvature
Deeply
cupped
NA2
NA2
Shallowly
cupped
Shallowly
cupped
Petal colour
White to pale
pink
Purple
Purple
Red
Orange
Sepal length x
width (mm)
7-10 x 3.5-4
8-10 x 2.5
8-9 x 3-3.5
10x4
6-7x2
Petal length x
width (mm)
6-8 x 3-7
6-8 x 3-4
9-10x5-9
8x5
7x3
Plant colour
Red
NA2
NA2
Green
Green
Geographic
location
Esperance
area
NA2
NA2
Perth area
Albany area
'The taxon D. microphylla var. macropetala and the autonym D. microphylla var. microphylla
were established in Diels ( 1906).
2This information is not provided in Diels (1906).
Table 1: A summary of variation recorded within the Drosera microphylla complex based on
descriptions in Diels ( 1 906; p. 119-121) and Lowrie (1987), and a comparison with plants from
the Esperance and Albany areas. Plant colour is taken from sun-exposed plants.
Acknowledgements:
I wish to thank Phill Mann for his assistance and company during recent field trips to the
Esperance area for further study of this taxon.
References:
Diels, L. 1906. Droseraceae. In A. Engler (Ed.) Das Pflanzenreich. 26: 1-137.
Lowrie, A. 1987. Carnivorous Plants of Australia: Volume 1 . University of Western Australia Press,
Perth.
Marchant, M.G. and George, A.S. 1982. Drosera. In: (A.S. George, Executive Editor) Flora of
Australia: Volume 8, pp. 9-64, 77. Australian Government Publishing Service, Canberra.
No boring plants!
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42
Carnivorous Plant Newsletter
Technical Refereed Contribution _
HELIAMPHORA EXAPPENDICULATA , A CLEARLY
DISTINCT SPECIES WITH UNIQUE CHARACTERISTICS
Joachim Nerz • AmdtstraBe 2* 71032 Boblingen • Germany
Andreas Wistuba • Mudauer Ring 227 • 68259 Mannheim • Germany
Keywords: new taxa: Heliamphora exappendiculata , Venezuela.
Received: 4 June 2003
Introduction
Due to its remoteness and inaccessibility, the flora of the Guyana Highlands, in the south of
Venezuela, remained nearly completely unexplored until as recently as only 50 years ago. Before
that time, only a few expeditions had reached the tepuis in the area. e.g. Roraima (Im Thurn 1 885),
Duida (Gleason 1931 ) or Auyan (Gleason 1939). The botanical collections from these expeditions
were amazing (Im Thurn 1886) — the flora is spectacular even in the extensive savannahs around
the tepuis (Schomburgk 1931 ).
In 1946, another pioneer in the exploration of the tepuis, Felix Cardona, explored Apacara-
tepui (a pail of Chimanta-tepui). On this trip he observed and collected material of an interesting
Heliamphora (Cardona 51648).
At approximately the same time, two groups of botanists started to explore this vast area more
systematically. One group was led by Julian A. Steyermark, who started his career at the Chicago
Natural History Museum. Later on, he became so fascinated in the botany of this extraordinary
region that he spent 25 years of his life in Venezuela. In 1984 he returned to the USA to the well-
known Missouri Botanic Garden. Never before or afterwards had anyone made so many botanical
collections as Steyermark. The other group was from the New York Botanical Garden and was led
by Bassett Maguire with the collaboration of John J. Wurdack. One of the largest tepuis. with a sur¬
face of 615 km2, the huge area of Chimanta-tepui was at that time unexplored. Steyermark and
Wurdack made independent expeditions to reach its top in 1953 (Steyermark 1955). Due to its
remoteness, Steyermark needed months to reach the top. Wurdack reached the top of Chimanta-
tepui on 29 January 1953. He collected material of this region’s interesting Heliamphora (Wurdack
34262; this material is used as the type for the species described in this paper) on the northwest
slopes of Churi-tepui, another part of the huge Chimanta-massif. In the same year, Bernardi orga¬
nized an expedition to reach the top of Aprada-lepui, an isolated tepui near and to the north of
Chimanta. Even though he failed in this goal, he was able to collect material (Bernardi 796) of the
same Heliamphora species at the base of this tepui. In 1955, Steyermark and Wurdack made anoth¬
er expedition to the Chimanta-massif, and collected yet additional material of this new
Heliamphora.
After these collections, this taxon was observed only very rarely. In 1978 it was described as
a variety of H. heterodoxa by Maguire (Maguire 1978). Between 1983 and 1986, Huber organized
several helicopter-expeditions to the Chimanta-massif and performed intensive botanical and geo¬
logical explorations. But in the book where he presented the results of these expeditions (Huber
1992) he never mentioned the distinctive plant. It seems that it was overlooked due to its habitats
which are both remote and difficult to access.
Heliamphora exappendiculata (Maguire & Steyermark) Nerz & Wistuba comb, et stat. nov.
(Memoirs of the New York Botanical Garden (Vol. 29, 1978): The Botany of the Guyana
Highland - Part X: p. 54) — ' Var heterodoxi simili sed amphoris sine appendicibus'
Holotype (NY): Venezuela, 29.01.1953, John .1. Wurdack 34262.
Rhizomes branching, plants often forming dense colonies. Plants growing in or below
Volume 35 June 2006 43
<S> IMAGED
KUNHAItDT-WlTHERBEE CHIMANTA MASSIF EXPEDITION, 1952-53
New York Botanical Garden — American Museum of Natural History
Estado Bolivar, Venezuela
No. 3^262 HOLOTYPE
Heliaraphora heterodoxa Steyermark
var. exappendiculata Maguire
det, Bassett Maguire, 1973
Petals pink, stamens yellow, pitcher appendage
slightly developed, frequent on escarpment face
in spray of falls, 1 km north of Camp 8 at lower
escarpment base, 2050 m elev.
Talus cloud forest, northwest slopes of Churi-tcpuf (Muru-tcpui).
John J. Wurdack January 29 ,'1953
Figure 1: Herbarium specimen 34262 (NY), holotype for Heliamphora exappendiculata.
Image provided courtesy of the New York Botanical Garden.
44
Carnivorous Plant Newsletter
vertical cliffs, with the rhizomes growing down, turning upwards and hearing pitchers. Below the
cliffs, the rhizomes grow upwards normally. Pitchers ventricose in the lower part, 12-25 cm long,
4-10 cm wide at the upper part. The outer surface of the pitchers loosely covered with simple
minute trichomes. The inner surface of the upper part of the pitcher densely covered with fine hairs.
The lid completely embedded in the apex of the pitcher, circular to ovate, 5-10 mm long, usually
pointed in the apex, approximately forming a small triangle at the inner side roughly. Outer surface
of the pitcher covered with short trichomes. Colour of the pitcher pale green, in sunny places it may
develop irregular red dots or completely red pitchers. Sometimes the pitchers are green in the lower
part and red in the upper part, especially on the inner pitcher surface. Lid usually red, slightly hairy
at the outer part. 1-3 flowers. Inflorescence about 40 cm long, peduncle 3-5 cm long, glabrous;
bracts 3-4 cm long, bearing rudimentary pitchers. Tepals 4, oblong-lanceolate, 3-5 cm long, 1 .5-2
cm wide, white to whitish-pink; 10 stamens in I series, filaments 7 mm long, anthers oblong, lance¬
olate, 8 mm long, 1 .2 mm wide. Ovary 3-celled, pubescent, style glabrous. Seed approximately 2
mm long, compressed, ovate, irregularly winged.
Specimens Examined
Churi-tepui (Muri-tepui), 2050 m, 29.01.1953, John J. Wurdack 34262, Holotype (NY);
Churi-tepui (Muri-tepui), 2050 m, 29.01.1953 Wurdack No. 34262, ISOTYPE (K); Churi-tepui
(Muri-tepui), 2250-3000 m, 26.01.1953 Wurdack No. 34236 (VEN); Churi-tepui, 2100-2200 m,
24.01.1953, Wurdack No. 34172 (VEN); Aprada-tepui, 1000-1100 nt. 18.08.1953, Bemardi No.
796 (VEN); Chimanta-massif, Central Section, 1925 m, 5.2.1955, Steyermark & Wurdack No. 441 ;
Torono-tepui, 1875-1950 m alt; 26.02.1955, Steyermark & Wurdack No. 1 135 (VEN); Apacapa-
tepui, 2125-2300 m, 13.04.1953 Steyermark No. 74888 (VEN, K); Apacara-tepui, 1900 m,
08.07.1946, Cardona No. 51648 (VEN); Amuri-tepui, 1850 m (+/-), 2-5.2.1983. Steyermark et al.
No. 128489 (VEN).
Geology and Geography
Heliamphora exappendiculata is apparently restricted to Chimanta-tepui and Aprada-tepui
(Brewer-Carias 1987), two sandstone-plateaus at the centre of the Gran Sabana. Chimanta is one
of the largest tepuis; it is characterized by many dissections, valleys and ravines at the plateau-area.
Aprada is smaller than Chimanta and is located close to it in the north-west. It consists of two
plateaus, both relatively fiat which are divided by a valley. Heliamphora exappendiculata seems to
be adapted to this high degree of vertical cliffs of Chimanta-tepui, where it grows with few other
plants. Due to the diversity of the surface of Chimanta-tepui, many different habitats can be found
on the plateau (including vast savannahs, shrubby forests, dry cliffs, wet cliffs and many rivers and
waterfalls). It is because of this great diversity of habitats, that three different species of
Heliamphora exist side by side at the top of Chimanta-tepui, Heliamphora exappendiculata, H.
pulchella and H. chimantensis. On Aprada only two species of Heliamphora have been recorded
so far, Heliamphora exappendiculata and H. pulchella. Here Heliamphora exappendiculata was
not found growing on cliff faces but rather in the shade on the bottom of small canyons of the
otherwise flat surface of the plateau.
Distribution
Heliamphora exappendiculata is restricted to the Chimanta-massif and Aprada-tepui. So far,
most collections have been made from the central and northern parts of these features. It also has
been recorded from the base and the top of Aprada-tepui. Heliamphora heterodoxa is only known
from the top of Ptari-tepui and from the adjacent areas in the Sierra de Lema. All specimens col¬
lected from Chimanta-tepui as H. heterodoxa (like e.g. Steyermark & Wurdack 374 and 375)
belong to the recently described new species H. chimantensis. From Auyan-tepui, Steyermark
(1984) described a distinct form as “ Heliamphora heterodoxa var. exappendiculata forma glabel¬
la (Steyermark).” The material was collected by Steyermark at the plateau of Auyan-tepui (J.A.
Steyermark 93712, VEN). Closer examination of this specimen, however, revealed that in fact it
Volume 35 June 2006
45
represents a highly etiolated H. minor plant. We also found material in Caracas labelled H. hetero-
doxa exappendiculata f. glabella , collected at the plateau of Aparaman-tepui. (Collected by Bruce
Holst at 22.03.1987 (Holst 3485) for the Missouri Botanical Garden herbarium and determined by
J.A. Steyermark.) It seems there was some confusion about the collected specimens of the
Aparaman-group, because members of the same taxon were also determined as H. heterodoxa
(Huber & Gorzula, 1 1.147, VEN), H. heterodoxa var. exappendiculata (Steyermark et al. 132045.
VEN). or remained unidentified (Holst et al. 2923, VEN). Indeed all specimens currently known
from the Aparaman-group belong to H. folliculata. These ambiguities can be resolved with current
understandings of the new species that have been subsequently described, i.e. H. exappendiculata
and H. folliculata. We hope, that in the future, the type-material for Heliamphora exappendiculata
f. glabella (Steyermark 93712) can be found again for further examinations.
Morphology and Distinguishing Characteristics
Heliamphora exappendiculata can easily be distinguished from other Heliamphora- species
because it is the only species where the lid is embedded in the surface of the pitcher. In contrast.
H. heterodoxa has one of the most prominent lids in the whole genus. The pitchers of H. exappen¬
diculata are infundibuliform. usually 12-15(25) cm long, whereas in H. heterodoxa the plant is in
all parts tubular and 20-30(40) cm long. The Bowers of H. exappendiculata are comparably large
with elongated tepals, usually 1-2 (lowered: in H. heterodoxa the flowers are comparably small
with the shortest tepals in the genus, usually 2-4(7) Bowers on each inflorescence. The ecological
differences are also obvious — H. exappendiculata grows at or below vertical, wet cliffs, while H.
heterodoxa grows exposed at the top of Ptari tepui or in open savannahs in the Sierra de Lema.
Relationships to Other Species
Heliamphora exappendiculata has no close similarities to H. heterodoxa. Heliamphora pul-
chella, which also grows at the Chimanta-massiv and on Aprada-tepui, shares with H. exappendic¬
ulata the short pitchers, but the other morphological characters of the pitchers, lid and flowers are
clearly different (see Table 1 ). Furthermore, the habitats of H. pulchella (open bogs and swamps)
are different from the habitats of H. exappendiculata. The other species from Chimanta-tepui that
grows near H. exappendiculata is H. chimantensis. The pitchers of these two species are clearly dif¬
ferent — those of H. chimantensis are tubular, elongated pitchers with a well developed lid. The
Bowers are also easily to distinguish; in H. chimantensis the tepals are shorter, lanceolate with
broad base, in H. exappendiculata the tepals are larger, and oblong-lanceolate.
Morphologically. H. exappendiculata shows most affinities to the recently described H. hisp-
ida (Wistuba & Nerz), which grows about 500 kilometers away at the isolated Neblina-plateau. It
has in common with H. hispida the short pitchers with a clearly infundibulate upper part, and also
the Bowers are similar with oblong-lanceolate tepals and 8-10 stamina. Furthermore, both species
grow in colonies. But there are clearly differentiating characteristics between these species, e.g. the
hairiness of the inner part of the pitcher, the absent lid in H. exappendiculata , and the different habi¬
tats, e.g. H. hispida never has been observed on vertical cliffs.
Etymology
The species epithet “ exappendiculata ” means "without appendage," and notes the unique
characteristic of this species.
Discussion
When Bassett Maguire and Julian A. Steyermark worked on the botany of the Guyana-shield
of Venezuela, they had to treat a huge amount of field and herbarium work. When they started their
expeditions, this extraordinarily rich flora was nearly unexplored, and even today many parts
remain to be explored. It took from 1 946, when H. exappendiculata was collected for the first time.
46
Carnivorous Plant Newsletter
H. exappendiculata
H. heterodoxa
H. pulchella
H. hispida
Pitcher
Dimensions
12-25 cm long
4-10 cm wide
1 5-40 cm long
5-6 cm wide
8-20 cm long
3-8 cm broad
15-25 cm long
5-8 cm wide
Shape
Ventricose in the
lower part,
infundibulate to
broad infundibulate
in the upper part
Infundibulate to
ventricose in the
lower third,
slightly
infundibulate in
the upper part,
expanded near
the mouth
Slightly ventri¬
cose in the lower
part, narrowly
expanded to
tubular in the
upper part
Slightly ventri¬
cose in the lower
part, broad
infundibulate in
the upper part
Lid
Dimensions
0.5-1 cm long
0.4-0. 8 cm wide
1-3.5 cm long
1-1.5 cm wide
0.5-1 cm long
0.3-0. 5 cm wide
1 - 1 .5 cm long
Shape
Round to ovate,
embedded in the
apical end of the
pitcher-surf ace, usu¬
ally pointed in the
apex
Strongly helmet¬
shaped, very con¬
stricted to stalked
at the base
Flattened to hel¬
met-shaped, only
slightly constrict¬
ed around the
base
Cordate, curved
Inflorescence
Dimensions
Peduncle 40 cm
long, pedicels 3-5
cm long
Peduncle 30-70
cm, pedicels 5-6
cm long
Peduncle 20-40
cm long, pedicels
1 2 cm long
Peduncle 100 cm
long, pedicels 12
cm long
Tepals
Oblong-lanceolate,
narrowing near the
base
Very broadly
lanceolate
Lanceolate,
broad base
Lanceolate,
broad base
Anthers/
Stamens
Dimensions
8 mm long
5.5-8mm long
4 mm long
3.5 mm long
Number
10
12
15
10
Table 1 : Comparison between H. exappendiculata. H. heterodoxa, H. pulchella and H. hispida
until 1 978 for this outstanding species to be described. It was first described only as a variety of H.
heterodoxa , perhaps because the authors — who were aware of its lack of a prominent lid — did not
appreciate the other divergent characters. Its description merely notes that "Var heterodoxi simili
sed amphoris sine appendicibus ” (“similar to var. heterodoxa , but pitchers without appendix”).
Further, it was mentioned that "in the var exappendiculata the appendage has become completely
or essentially lost, and the inner glandular area lowered to the apex of the leaf’. It seems that the
authors only concentrated on the most obvious characteristics — the lacking lid — but were not
aware of the other distinct characters of this species, which make it very different from H. hetero¬
doxa. Steyermark ( 1951 ) wrote of H. heterodoxa as being a variable species, but the specimen used
to justify this characterization (i.e. Steyermark 59766, with somewhat more infundibulate pitchers)
might have been collected in more shaded habitats. In actuality. H. heterodoxa is only a moderate¬
ly variable species. The specimens at the top of Ptari-tepui are narrower near the mouth and more
Volume 35 June 2006 47
colourful, compared to the usually greenish specimens that occur in the Gran Sabana or in shaded
habitats. In these areas they are somewhat more infundibuliform. Steyermark also felt that H. het-
erodoxa was a variable species because of two other collections, i.e. Steyermark & Wurdack 374,
and Steyermark & Wurdack 373. However, these specimens belong to the recently described
species H. chimantensis. When these incorrectly placed specimens are removed from inclusion
with H. heterodoxa , the species is seen as being not very variable at all.
We can conclude that H. exappendiculata is one of the most distinct and interesting
Heliamphora species, and one that can be identified clearly on the first sight.
Acknowledgements: An editor of this article (BR) wishes to thank staff of the New York Botanical
Garden for assisting with providing an image for this publication.
Literature
Brewer-Carias, C. 1987. The lost world of Venezuela and its vegetation. Caracas, Venezuela.
Gleason, H.A. 1931. Botanical Results of the Tyler-Duida expedition. Bull, of the Torry Club. 58:
277-506.
Gleason. H.A. 1939. The flora of Mount Auyantepui. Brittonia 3: 164.
Huber. O. 1992. El macizo del Chimanta. Oscar Todtmann Editores.
Im Thum, E.F. 1885. The first ascent of Roraima. Timehri. 4: 1-48.
1m Thum, E.F. 1 886. Notes of the plants observed during the Roraima expedition of 1 884. Timehri
5: 147-223.
Maguire, B. 1978. Sarraeeniaceae ( Heliamphora ) in The Botany of the Guyana Highland Part-X.
Mem. of the New York Bot. Garden 29: 36-61.
Nerz, J. and Wistuba. A. 2000. Heliamphora hispida (Sarraeeniaceae), a New Species from Cerro
Neblina, Brazil-Venezuela. Camiv. PI. Newslett. 29: 37-41.
Schomburgk, R.H. 1931. Travels in Guiana and on the Orinoco. Leipzig. English translation by
Walter E. Roth, Georgetown. British Guiana.
Steyermark, J.A. 1951. Sarraeeniaceae in Botanical Exploration in Venezuela - I. Fieldiana Bot.
' Vol. 28(1): 239-241.
Steyermark. J.A. 1955. Attack on Chimanta. Nat. Hist. Vol. 64(9): 482-488.
Steyermark. J.A. 1984. Realignment of the Genus Heliamphora. Annals of the Missouri Botanical
Garden 71: 302-312.
Wistuba, A., Harbarth , P, and Carow, T. 2001. Heliamphora folliculata, a new species of
Heliamphora (Sarraeeniaceae) from the ‘Los Testigos' Table Mountains in the South of
Venezuela. Camiv. PI. Newslett. 30: 120-125.
Wistuba, A.. Carow, T., and Harbarth, P. 2002. Heliamphora chimantensis, a new species of
Heliamphora (Sarraeeniaceae) from the 'Macizo de Chimanta’ in the South of Venezuela.
Camiv. PI. Newslett. 31: 78-82.
Wistuba, A., Carow. T.. Harbarth , P, and Nerz, J. 2005, Heliamphora pulchella, eine neue mit
Heliamphora minor (Sarraeeniaceae) verwandte An aus der Chimanta Region in Venezuela.
Das Taublatt. 53/3: 42-50.
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48
Carnivorous Plant Newsletter
Technical Refereed Contribution _
Naphthoquinones Content of Some Sundews (Drosera L)
Jozef Kovacik • Miroslav RepcAk • Department of Botany • Institute of Biology and Ecology
• Faculty of Science • P. J. Safarik University • Manesova 23 • 041 67 Kosice • Slovak Republic
• jozkovacik@yahoo.com
Keywords: chemistry: Drosera.
Introduction
Naphthoquinones are distributed within eight families of (lowering plants and are known
also in fungi, lichens, and in some spiders. These compounds are characteristic for Droseraceae;
they were detected in genera Dionaea , Aldrovanda and Drosophyllum (Culham & Gornall,
1994). Plumbagin (5-hydroxy-2-methy 1-1, 4-naphthoquinone) and 7-methyljuglone (5-hydroxy-
7-methyl- 1,4-naphthoquinone) are two main naphthoquinones of the genus Drosera (Bonnet el
al., 1984). Plumbagin was also detected in the genus Nepenthes and in the genera
Triphyophyllum and Dioncophyllum of Dioncophyllaceae. Within angiosperms, 7-methyljuglone
is the predominant quinone in Ebenaceae and plumbagin in Plumbaginaceae and Iridaceae
(Culham & Gornall. 1994). Aglycones droserone (3,5-dihydroxy-2-methyl-l ,4-naphthoquinone)
and hydroxydroserone (3,5,8-trihydroxy-2-methyl-l,4-naphthoquinone) are present in minor
amounts (Samaj et al., 1999). Other known minor naphthoquinones are diomuscione and
diomuscipulone, identified in Dionaea muscipula (Miyoshi el al.. 1984). In sundews these com¬
pounds probably act as allelopathic, molluscidal, antimicrobial, and antifeedant chemicals
(Samaj et al., 1999).
The taxonomic significance of naphthoquinones within Droseraceae was studied by
Culham & Gornall (1994). They found that their occurrences within various species were not
useful taxonomic tools because closely related taxa (e.g., subspecies of Drosera stolonifera) did
not have the same naphthoquinone. Mainly D. rotnndifolia was studied for naphthoquinones con¬
tent due to its pharmacological use (e.g., Galambosi et al., 2000; Repcak et al., 2000). The 7-
methyljuglone content in D. rotnndifolia and D. anglica in dependence on ontogenetic phases of
the leaf were studied by Repcak et al. (2000). The highest content was found in young and adult
green leaves, while it was lower in older leaves.
In this paper, the content of 7-methyljuglone and plumbagin was studied in several non-
European Drosera species.
Figure 1: From left to right, 7-methyljuglone, plumbagin, droserone, hydroxydroserone.
Materials and Methods
Plants were cultivated in a greenhouse of the Botanical Garden of P. J. Safarik University at
Kosice (Slovak Republic) in pots with peat and were watered with distilled water. For 7-
methyljuglone and plumbagin estimation, we harvested fully developed leaves of comparable
age from six different plants of each species. Fresh material was homogenized and extracted with
Volume 35 June 2006
49
400-'
Figure 2: HPLC chromatograms of 7-methyljuglone and plumbagin in extracts from
fresh leaves of Drosera capensis, D. binata, and their mixture.
benzene, evaporated to dryness, dissolved in methanol, and analysed by an isocratic HPLC sys¬
tem. Compounds identity were detected according to their TR (retention time). UV-V1S spectrum
was measured during the analysis to confirm the identity of naphthoquinones. Juglone (Fluka)
was used as an internal standard. HPLC conditions: column Biospher SI C 18, 7 pm (Tessek.
Prague, Czech Rep.) - 3.3 x 150 mm. flow rate 0.5 ml min F The mobile phase was 50 % ace¬
tonitrile (Merck). The detection was performed at 421 nm.
Results and Discussion
We detected 7-methyljuglone in nine out of eleven analysed species and plumbagin in D.
binata and D. prolifera. Very different minimal and maximal values in each species shown in
Table 1 could reflect different leaf age and support the well-known fact that naphthoquinone con¬
tent is negatively correlated with the age of tissue (Repcak & Galambosi, 1994). It is difficult to
collect the leaves from the same position on the stem, e.g., in sundews forming leaf rosettes. In
spite of this fact, on the basis of naphthoquinone content, we can divide the analysed species into
three groups.
The first group, with the highest content of these compounds, is represented by D. collinsi-
ae. D. capensis , and D. binata. More extensive quantitative data were published by Caniato et
al. (1989). In aboveground parts in D. binata. they found a mean plumbagin content 0.23-0.31
% (per unit fresh weight). For this species, we found 0.31 % of plumbagin in fresh leaves. In
aboveground parts in D. capensis. they found 0.10-0.1 1 % (FW) of 7-methyljuglone. For this
species, we found a value of 0.36 % in fresh leaves. Note that in the sepals and the pistil of D.
capensis. we detected 7-methyljuglone and, in the same parts of D. binata flower, we detected
plumbagin.
The second group (D. communis. D. venusta. D. admirabilis, and D. spathulata ) had a
medium content of the analysed compounds but 3-5 times smaller when compared to the first
group. No quantitative data referring the content of 7-methyljuglone in some species of this
group have been found in the literature.
The third group with low content of the analysed compounds includes D. burkeana, D. ade-
lae. and D. hamiltonii. They contain approximately 10-times less 7-methyljuglone in compari¬
son with the first group. The position of D. prolifera is somewhere in between the first and sec¬
ond group. The variance of values is narrow what can be satisfactorily explained by the shape of
leaf rosette when it is easy to determine the leaves of the same age on different plants.
As reported in the literature (Culham & Gornall, 1994), naphthoquinones are present in all
four genera of the Droseraceae. The plumbagin content in Aldrovanda vesiculosa collected from
50 Carnivorous Plant Newsletter
7-methyljuglone
X±SD
min.
max.
Drosera collinsiae
46.0±12.6 a
24.7
61.6
Drosera capensis
40.8±7.39 ab
33.0
50.5
Drosera communis
9. 14±5.05 cd
2.63
14.9
Drosera venusta
8.54±3.25 cd
7.16
11.6
Drosera admirabilis
8.19+2.99 cd
4.63
13.0
Drosera spathulata
7.88±5.42 cd
3.18
16.1
Drosera burkeana
4.93±3.32 d
1.26
8.86
Drosera adelae
4.03±2.14 d
1.80
6.69
Drosera hamiltonii
3.83±2.34 d
1.26
7.34
Plumbagin
Drosera binata
30.3± 8.70 b
22.3
46.2
Drosera prolifera
18.7+1.66 c
16.9
21.5
Table 1 : Naphthoquinones content in Drosera species (mg g' * of dry weight). Values in the ver¬
tical column followed by the same letter do not differ significantly at P<0.05 by Tukey’s pair¬
wise comparison; n=6; X, mean value; SD, standard deviation; min./max., minimal/maximal
detected value.
different sites or originating from different continents is similar to the content of naphtho¬
quinones in the first group with the highest content of these compounds (for comparison see
another paper in this issue of Carnivorous Plant Newsletter; Adamec, et al. 2006).
References
Adamec, L., Gastinel, L., Schlauer, J. 2006. Plumbagin content in Aldrovanda vesiculosa shoots.
Carniv. PI. Newslett. 35: 52-55.
Bonnet, M., M. Coumans, M. Hoftnger, J.L. Ramaut, and Th. Gaspar. 1984. High-performance
gas chromatography of 1,4-naphthoquinones from Droseraceae. Chromatografia, 18: 62 1 -
622.
Caniato, R.. R. Filippini, and E.M. Cappelletti. 1989. Naphthoquinone contents of cultivated
Drosera species Drosera binata , Drosera binata var. dichotoma and Drosera capensis. Int.
J. Crude Drug., 27: 129-136.
Culham A., and R.J. Gornall. 1994. The taxonomic significance of naphthoquinones in the
Droseraceae, Biochem. Syst. and Ecol., 22: 507-515.
Galambosi, B., Zs. Galambosi, and M. Repcak. 2000. Growth, yield and secondary metabolites
production of Drosera species cultivated in peat beds in Finland. SOU, 51: 47-57.
Miyoshi. E., Y. Shizuri, and S. Yamamura. 1984. Isolation and structures of diomuscione and
diomuscipulone from Dionaea muscipula. Phytochem., 23: 2385-2387.
Repcak, M., and B. Galambosi. 1994. Ontogenetic aspects of 7-methyljuglone accumulation in
some Drosera L. species. Biol. Plant., 36: S232.
Repcak, M., B. Galambosi, and N. Takkunen. 2000. The production of 7-methyljuglone,
quercetin and kaempferol by Drosera anglica and Drosera rotundifolia. Biologia,
Bratislava, 55: 429-433.
Samaj, J., A. Blehova, M. Repcak, M. Ovecka. and M. Bobak. 1999. VII. Drosera species (sun¬
dew): in vitro culture and the production of plumbagin and other secondary metabolites. In:
Biotechnology in agriculture and forestry, 43. Med. and aromat. plants XI, ed. Bajaj I. P. S.,
Springer- Verlag, Berlin, 105-135.
Volume 35 June 2006
51
Technical Refereed Contribution
Plumbagin Content in Aldrovanda vesiculosa Shoots
Lubomir Adamec • Institute of Botany • Dukelska 135 • CZ-379 82 Trebon • Czech Republic •
adamec@butbn.cas.cz
Louis Gastinel • University of Limoges • Faculty of Sciences and Technique • EA 3176 Plant
Glycobiology and Biotechnology • 123 Avenue A. Thomas • F-87060 Limoges • France •
lgastinel@unilim.fr
Jan Schlauer • University of Tibingen • ZMBP, Auf der Morgenstelle 5 • 72076 Tubingen •
Germany • jan@carnivorousplants.org
Keywords: chemistry: Aldrovanda vesiculosa.
Introduction
Plumbagin is a yellow naphthoquinone (for its chemical structure see Kovacik & Repcak in
another paper of this issue) typically occurring in all genera ( Drosera . Dionaea. Aldrovanda) of
Droseraceae and widespread within the order Nepenthales (Bonnet et al., 1984; Juniper et al.,
1989; Culham & Gomall, 1994, Tokunaga et al.. 2004; Schlauer, 2005). Although the presence
of plumbagin or its topoisomer, 7-methyljuglone, in Droseraceae organs can very simply be
proved by spontaneous sublimation of these substances and staining plastic materials and organ¬
ic solvents yellow, both the correctness of analytical procedures for quantitative determination of
plumbagin and mainly its physiological functions in (carnivorous) plants are still a matter of dis¬
cussion. Moreover, plumbagin and its derivatives have long been under the interest of pharma¬
cologists for their antimicrobial effects. Tokunaga et al. (2004) have recently demonstrated
strong cytotoxicity of plumbagin against cancer cells and also its antifeedant effect against insect
larvae.
In Drosera and Dionaea leaves, plumbagin content usually ranges between 2-3% of dry
weight (DW; Tokunaga et al.. 2004; Kovacik & Repcak, 2006). In the rootless aquatic carnivo¬
rous plant Aldrovanda vesiculosa L., plumbagin content has never been determined quantita¬
tively. As sun-adapted Aldrovanda plants of green temperate European and Asian populations are
typically yellow-green and completely lack anthocyanins, a possibly high plumbagin content in
this species could be associated with a photoprotective role. The aim of this paper was to inves¬
tigate plumbagin contents among all available Aldrovanda populations, compare two methods of
quantitative plumbagin determination in Aldrovanda. and to determine plumbagin content in
Aldrovanda plants of different populations or having been grown at different irradiances. Thus,
the possible photoprotective role of plumbagin was tested.
Materials and Methods
Plant material of Aldrovanda for plumbagin content determination was collected from both
field sites and outdoor or indoor cultures on 20 August 2004. Adult plants originating from E
Poland were collected from artificial sites Ptaci blato 1st pool, and Karstejn fen lake in the
Trebon region, S Bohemia. Czech Republic (plants were introduced to these sites, see Adamec
& Lev. 1999; Adamec, 2005). Plants at Ptaci blato grew in slight shade, while those at Karstejn
were collected from three microsites differing greatly among each other in the level of shading
by emergent reed vegetation; sun-adapted plants without any shading, shade-adapted plants, and
plants growing in very deep shade. The latter plants grew in dense reed stand, evidently at the
light minimum threshold for their growth. They were dark green, very short (only 4-5 cm), and
weak. Polish plants were also collected from two outdoor plastic cultivation containers in which
they had grown in slight shade (Adamec, 1997). Red plants of three Australian tropical popula¬
tions from outdoor cultivations (N Australia near Darwin, NT; NW Australia from Kimberley,
52
Carnivorous Plant Newsletter
NT) or an indoor cultivation (N Australia near Katherine, NT) were also collected. They were
grown in slight shade (Adamec, 1999). Ripe turions of Polish plants were collected from Ptaci
blato and two outdoor containers on 23 October 2004.
For a qualitative test of plumbagin presence, plant material of fourteen Aldrovanda popula¬
tions was collected from small aquaria in an outdoor collection of plants maintained by one of
the authors (LA). The original provenances of these plants were: S and N Russia, N Ukraine,
Lithuania, E and W Poland, SW Hungary, S Germany (now Switzerland), Japan (near Tokyo), N
Australia (Katherine and Darwin), NW, SE, and SW Australia (for most of them see Maldonado
San Martin et al., 2003).
Collected plants were thoroughly washed and shoots were divided into two segments, apex
with six adult leaf whorls (denoted as “apex”) and subsequent six leaf whorls (7th- 12th whorls;
denoted as "base”). One apical or basal shoot segment (fresh weight 28-145 mg, DW 2.3-12 mg)
or one turion (DW ca. 4 mg) was used for one determination. Segments or turions were blotted
dry and extracted three times with 1 ml diethyl-ether for about 20 min. before DW of the bio¬
mass was estimated. Pooled extracts were alkalized by adding 1 ml 0. 1 M NaOH and thorough¬
ly shaken for a few sec. As a result of alkalization, the extract turn orange-red ( i.e., bathochromic
effect) and plumbagin was dissolved in the aqueous phase. The aqueous phase was diluted ten
times with distilled water and optical density measured against blank sample at 400 nm.
Plumbagin standard (practical grade P7262, Sigma) was used to obtain calibration curve in the
same way. Results are expressed in % DW. Although this method of plumbagin extraction and
determination is very simple and fast some objections may arise as to the specificity of this deter¬
mination. Four parallel plants of each variant were dried shortly at 40°C, wrapped to an alu¬
minum foil, and sent by post to the Laboratory of L. Gastinel (Univ. Limoges, France) for HPLC
method of plumbagin determination. After they were overwintered in a refrigerator at 3 C, liv¬
ing turions from Ptaci blato were also sent by post to the Laboratory of L. Gastinel on 17 March.
Here, 3 to 20 mg DW of Aldrovanda shoots were extracted in glass tubes with 2 ml of
diethyl-ether three times for 30 min. The pooled extracts were allowed to dry overnight by evap¬
oration. The extracts were then resolubilised with I ml of pure methanol in the presence of 0. 1 %
formic acid (FA). Twenty-five |il of the samples were injected on an HPLC column (Nucleosil
250 x 4.6 mm 300 A C18 5p with a guard column at a 0.5 ml min-1 flow rate. Elution of sub¬
stances absorbing at 400 nm (naphthoquinone ring) was followed by using a 45-min linear gra¬
dient from 50% methanol: 50% water with 0.1% FA to 80% methanol: water with 0.1% FA.
Quantitative calibration of plumbagin was performed using the plumbagin standard (see above)
dissolved in pure methanol with 0.1% FA and injected in the HPLC at different volumes.
MS spectra of the 400 nm absorbing material purified from HPLC gradient (retention time
around 21 minutes) were recorded after the injection in the Turboion ESI spray coupled with Q-
trap mass analyser (Applied Biosystems) with parameters, ion spray voltage 5000 V, decluster¬
ing potential 30 V, How rate 40 pi min f m/z peaks were recorded in the range 50 to 500 amu
with the EMS protocol from Analyst 1.4 (Applied Biosystems).
Plumbagin presence was tested using a thin-layer chromatography. One shoot apex of each
population, 6-8 mm long, was blotted dry and extracted with 0.25 ml diethyl-ether for 15 min.
The extract (20 pi) was dropped on a TLC silica plate F60 and developed first in methanol (to
focus the front) and then in toluene for about 15 min. Leaf extract of a non-flowering plant of
Drosera intermedia was co-chromatographed as standard for plumbagin detection.
Results and Discussion
The presence of plumbagin was clearly determined in all fourteen world populations of
Aldrovanda but 7-methyljuglone was not detected. Comparing the two methods of plumbagin
content determination, direct colorimetric determination (see Table 1) and that using HPLC (see
Table 2), there is a much better agreement of the results for apical than for basal segments. Direct
colorimetric determination in the same material led to shoot contents greater by 0-90% in apices.
Volume 35 June 2006
53
while 2-5 times greater in bases, than using HPLC. There are some possible reasons for this dif¬
ference. First, the diethyl-ether extract at the direct colorimetric determination might contain also
some other ether-extractable substances absorbing at 400 nm, although no such contaminant has
been identified so far. Second, due to sublimation of plumbagin, a good deal of this volatile sub¬
stance could be lost during drying or posting the plant material or solubilized in the plastic phase
of the aluminum foil. It is conceivable this loss affected particularly the basal stem segments and
Australian plants with more opened leaves or lower biomass, in which the differences between
the two methods were more pronounced than in the apices or turions.
Overall, plumbagin content in apical segments ranged between 1.2-4. 8% DW and that in
basal ones between 0. 4-5.0% DW (see Tables I. 2). The mean plumbagin content of about 2-3%
DW in Aldrovanda apical shoot segments corresponds to that found in two Drosera species ( 1 .9-
3.0% DW; Kovacik & Repcak. 2006) or in Dionaea muscipula (about 3 % DW; Tokunaga et al.
2004). Thus, plumbagin content in Aldrovanda shoot is not greater than in other genera of
Droseraceae. However, with either assay method, plumbagin content was evidently greater in
apical than basal shoot segments in Polish plants but this difference was not too distinct in dark-
red Australian plants (see Tables 1, 2). This gradient in plumbagin content in Aldrovanda shoot
segments of different age is in a good agreement with Repcak et al. (2000) who found a decrease
of 7-methyljuglone content in older leaves of Drosera spp. Obviously, plumbagin is released
from old and ageing tissues of Aldrovanda to ambient medium (Adarnec, unpubl.). In
Aldrovanda , plumbagin content in shoot apices was similar to that in turions (Tables 1, 2). The
data on plumbagin content do not support the hypothesis that this substance has a specific and
light-regulated photoprotective role in Aldrovanda as the content was the same in both sun-and
shade-adapted plants. Nevertheless, due to optical properties of plumbagin (absorption of blue
light and UV), it is possible to assume certain role in photoprotection. These results support the
view that the main and primary role of plumbagin in Droseraceae is antifeedant and antimicro¬
bial, thus protecting the plants against herbivores and microbial parasites (Tokunaga et al. 2004).
Origin and light exposure of plant material
Plant A
Plant B
Apex
Base
Apex
Base
Shoot segments
E Poland: Ptaci blato 1st pool, slight shade
3.34
1.66
3.13
1.67
E Poland: Karstejn fen lake, sun-adapted
2.75
2.00
2.54
1.68
E Poland: Karstejn fen lake, shade-adapted
2.18
1.70
2.37
1.47
E Poland: Karstejn fen lake, very deep shade
2.31
1.46
3.04
2.49
E Poland: culture I; small container, slight shade
3.78
2.78
3.64
2.54
N Australia, Katherine: indoor aquarium, slight shade
4.55
3.41
5.95
3.93
Turions
Turion A
Turion B
E Poland: Ptaci blato 1st pool, slight shade
2.68
3.21
E Poland: culture I; small container, shade
2.69
2.39
E Poland: culture 11: big container, shade
3.50
3.68
Table 1 : Content of plumbagin (in % of DW) in apical and basal shoot segments of different
strains and in ripe turions of Aldrovanda vesiculosa. Apical shoot segments contained shoot
apex + the first six adult leaf whorls, while the basal ones the subsequent six leaf whorls (i.e.,
the 7th- 12th ones). Results for two different plants or turions A and B are based on measure¬
ment of optical density at 400 nm of alkalized diethyl-ether extract.
54
Carnivorous Plant Newsletter
Origin and light exposure of plant material
Plumb, content (% DW)
Apex
Base
Ptaci blato
2.2±0.9a
0.4±0.07b
Karstejn: sun-adapted
2. I±0.9a
0.5±0.05b
Karstejn: shade-adapted
2.4±0.4a
0.7±0. 1 b
Karstejn: deep shade-adapted
1.2±0.4
—
Culture I: small container
2.0±0.7a
0.5±0.1b
NW Australian: outdoors
1.2±0.2a
0.5±0.06b
Turions-Ptaci blato: 1st pool
2.4+0. 5
Table 2: Plumbagin content determined in diethyl-ether-extracted dry Aldrovanda biomass
using HPLC. For apical and basal segments, see Table 1 . Except for Australian plants, the oth¬
ers were from E Poland. Means+SD of 3-4 independent analyses are shown. The different let¬
ters within the same raw denote statistically significant difference at P<0.05 ( t-test).
References
Adamec, L. 1997. How to grow Aldrovanda vesiculosa outdoors. Carniv. Pi. Newslett., 26: 85-88.
Adamec, L. 1999. The biology and cultivation of red Australian Aldrovanda vesiculosa. Carniv. PI.
Newslett., 28: 128-132.
Adamec, L., and Lev, J. 1999. The introduction of the aquatic carnivorous plant Aldrovanda vesicu¬
losa to new potential sites in the Czech Republic: A five-year investigation. Folia Geobot., 34:
299-305.
Adamec, L. 2005. Ten years after the introduction of Aldrovanda vesiculosa to the Czech Republic.
Acta Bot. Gall., submitted.
Bonnet. M., Coumans, M., Hofinger, M.. Ramaut, J.L., and Gaspar, Th. 1984. High-performance
gas chromatography of 1,4-naphthoquinones from Droseraceae. Chromatografia, 18: 62 1 -
622.
Culham, A., and Gomall, R.J. 1994. The taxonomic significance of naphthoquinones in the
Droseraceae. Biochem. Syst. Ecol., 22: 507-515.
Juniper, B.E., Robins, and R.J., Joel, D.M. 1989. The Carnivorous Plants. Academic. London.
Kovacik, J., and Repcak, M. 2006. Naphthoquinones content of some sundews (Drosera L. ).
Carniv. PL Newslett., 35: 49-51.
Maldonado San Martin, A.P.. Adamec, L., Suda, J., Mes, T.H.M., and Storchova, H. 2003. Genetic
variation within the endangered species Aldrovanda vesiculosa (Droseraceae) as revealed by
RAPD analysis. Aquat. Bot., 75: 159-172.
Repcak, M., Galambosi, B., and Takkunen, N. 2000. The production of 7-methyIjuglone, quercetin
and kaempferol by Drosera anglica and Drosera rotundifolia. Biologia (Bratislava), 55: 429-
433.
Schlauer, J. 2005. Carnivorous Plant Chemistry, Acta Bot. Gallica, in press.
Tokunaga, T„ Dohmura, A., Takada, N„ and Ueda. M. 2004. Cytotoxic antifeedant from Dionaea
muscipula Ellis: A defensive mechanism of carnivorous plants against predators. Bull. Chem.
Soc. Jap., 77: 537-541.
Looking Back: CPN 25 years ago
Previous CPN coeditor and ICPS cofounder Don Schnell published a now-classic review of
all the infraspecific Sarrcicenia purpurea taxa then known, including a variation that resulted
because of its growing in marl fens. After twenty-five years, this treatment would be modified by
the addition of Sarracenia purpurea subsp. venosa var. montana , Sarracenia purpurea subsp.
venosa var. burkei f. luteola. perhaps a nod to the veinless expression of Sarracenia purpurea
subsp. purpurea. Of course a modern author would also have to address the issue of whether to
accept the Sarracenia rosea species as having merit.
Volume 35 June 2006
55
International Correspondent
Book Review
Correa A., M.D., and dos Santos Silva, T.R. 2005. Drosera (Droseraceae): Flora Neotropica
Monograph 96. 0-89327-463-1, 72 p., black and white figures and maps. Paperback,
US$25. 00+shipping. Order from www.nybg.org or 718-817-8721.
Reviewed by Fernando Rivadavia
This 72-page publication (in Spanish) on Droseraceae is a new addition to the Flora
Neotropica series, published by the New York Botanical Garden. The authors M.D. Correa A. &
T.R.S. Silva have published a few works on Droseraceae in Latin America over the past 30 years,
including the recent D. peruensis T.Silva & M.D. Correa.
After a brief introduction and taxonomic history of the genus Drosera in the New World
tropics, the authors describe general morphology of native species, and present interesting scan¬
ning electron microscope images of seeds and glands on leaves. The sections that follow cover
cytology, palynology (with more SEM images), phytochemistry, infrageneric classification, geo¬
graphic distribution (with numerous maps covering every species), floral biology and seed dis¬
persal, ecology, conservation, human uses, and finally the taxonomic treatment which takes up
nearly forty pages with detailed descriptions of each species, including morphology & distribu¬
tion.
The first impression is that this is a very impressive work, probably one of the largest and
most complete taxonomic publications on Drosera since Diels’ monograph in 1906 or Allen
Lowrie's three volume treatment of the Carnivorous Plants of Australia. The authors should at
least be praised for the effort of bringing together so much information spread throughout dozens
of publications spanning mostly the last century.
Unfortunately, only a slightly less superficial reading of this publication will bring to light
a disheartening endless series of oversights and errors. These range from numerous misspellings,
maps with dots in wrong places, and contradictory statements (sometimes even in succeeding
paragraphs) to more serious mistakes such as quotes from important publications (including
Juniper, Robins & Joel’s "The Carnivorous Plants” and Takahashi & Sohma’s "Pollen
Morphology of the Droseraceae and its Related Taxa”) which were wrongly interpreted by the
authors, possibly due to a poor grasp of English.
Knowledgeable botanists may identify further signs of worry, such as the less-than-precise
botanical drawings, the often vague morphological terminology, the non-inclusion of important
herbarium collections from the past decade (especially those at the University of Sao Paulo,
Brazil), the non-citation of more recent molecular phylogenetic data, and the obvious failure to
even attempt addressing the taxonomic difficulties posed by the more complicated species com¬
plexes. The odd assortment of taxa dumped under D. montana A.Saint-Hilaire, with no recogni¬
tion even of varieties, is absolutely indefensible.
The reader will hopefully remember that this work is not the final say in the matter of the
Droseraceae of the New World tropics. Unfortunately the recently published D. camporupestris
F.Rivadavia, D. grantsaui F.Rivadavia, D. tentaculata F.Rivadavia, and D. viridis F.Rivadavia
were not mentioned in this publication. New species will certainly continue to be described from
this biologically wealthy area and new taxonomic revisions will be made.
The overall sum of problems detected in this publication is simply inexcusable for two main
reasons. First of all, most of the errors could and should have been pointed out by careful review¬
ers. Second, the authors seemingly did not realize the importance of this publication, that any
and all errors published, as well as inadequate taxonomical treatments, will be propagated and
multiplied — since this will be a reference to numerous other authors for at least decades to come.
Unfortunately. Flora Neotropica Volume 96 illustrates a failure in the process of authoring,
editing, and reviewing a scientific work, and its legacy will be with us for a long time. I have not
reviewed the other volumes in the series and so cannot say if this problem is limited to the treat¬
ment of the Droseraceae.
56
Carnivorous Plant Newsletter
Literature Reviews
Akhriadi, P., Hemawali, and Tamin, R. A. 2004. A new species of Nepenthes (Nepenthaceae) from
Sumatra. Nuytsia 15: 355-393.
The authors identify Nepenthes rigidifolia as a new highland species from north Sumatra. This is
apparently the same entity as discovered by C. Lee. and described as " Nepenthes species A” in Clarke
(2001, Nepenthes of Sumatra and peninsular Malaysia); it has also been given other trade names by
nurseries. Nepenthes rigidifolia has affinities to N. hang so, N. ovata , and N. spectabilis , and the vari¬
ous differences are described in the paper — there are no single “spot characteristics” that easily identi¬
fy this species.
It is always a pleasure to see floristic works being performed by regional botanists. However, I
encourage authors, especially those writing in a second language, to seek assistance in drafting their
papers — the grammar in this work is sometimes confusing and detracts from the clarity of the overall
findings. (BR)
Aim, T. 2005. Pinguicula vulgaris (Lentibulariaceae) and its uses in Norway. Sida. 21: 2249-2274.
This comprehensive review of how this plant was (and still is!) utilized in Norway is a treasure-
trove for those seeking evidence of strangeness in humanity. For examples, Pinguicula vulgaris has
been given many common names, my favorites being “gnome-bite grass,” “slime-grass”, "old woman's
mouth,” and quite inexplicably “thickening old man.” Its connection to making tettemelk is discussed,
and a photograph is presented of such a viscous milk (Tjukkmjplk) commercially available in Norway.
Furthermore, we learn that P. vulgaris is useful to treat soreness in reindeer teats, and can be used to
calm down cows that have mated. Furthermore, it has some minor magical properties: if laid under a
girl’s pillow, she would dream of their coming husband. And that’s not all. . . ( BR )
Berry, P.E., Holst, B.K., Yatskievych, K. (eds.), 1998-2005. Flora of the Venezuelan Guayana. Vols 4:
697-703, 5: 782-803, 9: 138-144.
With the publication of Volume 9, the treatment of the last of the carnivorous monocots of the
tepuis is complete. In total, fourteen Drusera , seven Genlisea, forty-six Utricularia , and ten
Heliamphora are discussed with keys, line drawings, and short descriptions. This is an excellent
resource for those interested in the species from this region. Of course, new species have been described
that are not described in these treatments; such new species are noted in the literature reviews of
Carnivorous Plant Newsletter. (BR)
Carow, T. 2005. Fleischfressende Ptlanzen. Kosmos Garten. ISBN 3-440-10176-2 (in German)
Master carnivorous plant photographer Thomas Carow has written a small (48 page), illustrated
book on carnivorous plants. An introductory work, it discusses horticulture of carnivorous plants in the
home and garden, minibogs, and terraria. Short treatments are given of the carnivorous genera. Carow’s
photographs, of course, are delightful! Interested customers should contact German colleagues to coor¬
dinate a purchase. If nothing else, copies can be bought directly from the author (carow@t-online.de)
for approximately US$10, including shipping. (BR)
Degtjareva, G., Casper, J., Hellwig, F. & Sokoloff, D. 2004. Seed morphology in the genus Pinguicula
(Lentibulariaceae) and its relation to taxonomy and phylogeny. Botan. Jahrb. Syst. 125: 431-452.
The title is slightly misleading, as the species studied are predominantly Eurasian, and only one
(P. moranensis) from Mexico. Although the authors are convinced micromorphological features of the
seed surface are significant at sectional, specific or infraspecific level, the most striking characteristic
of the studied seeds is their similarity. Particularly in comparison with the huge structural diversity
Volume 35 June 2006
57
found in the seeds of the closely related genus Utricularia, the seeds of Pingaicula are best described
as essentially uniform. The members of P. sect. Pinguicula (to which most of the studied species
belong) share the presence of furrows between the outer parts of the anticlinal walls of adjacent
exotesta cells and an always monocotyledonous embryo. (JS)
Garrido, B., Harnpe, A.. Maranon, T., and Arroyo, J. 2003. Regional differences in land use affect
population performance of the threatened insectivorous plant Drosophyllum lusitanicum
(Droseracae). Diversity and Distributions. 9: 335-350.
The authors surveyed Drosophyllum sites and assigned four age classes to the plants
observed. This allowed them to infer how well new plants were being recruited, and to under¬
stand more about the status of the populations in the wild. The five sites studied in Morocco were
damaged from heavy grazing, although the plant's range in this country was probably never
extensive. The twenty Spanish sites included the places most likely to retain Drosophyllum in the
future, since they included populations in the Los Alcornocales Natural Park. The seven
Portuguese sites are highly impacted by development of housing and infrastructure, and also
plantations. Seedling recruitment at such sites is poor, and Drosophyllum is likely to go extinct
at these sites. (BR)
Lowrie, A. 2005. A taxonomic revision of Drosera section Stolonifera (Droseraceae) from south¬
west Western Australia. Nuytsia 15: 355-393.
All taxa of section Stolonifera (“fan-leaved" sundews, of which some have peltate and not
fan-shaped leaf blades) were already featured in Carnivorous Plants of Australia, Vol. 1, by the
same author (Univ. of W. Australia Press. 1987, ISBN 0 85564 254 8), from which the drawings
have been reproduced. This time the species descriptions are more detailed than in the book: syn¬
onyms, basionyms and types are cited, and lists of examined specimens are added (but maps and
colour photographs are missing). SEM photomicrographs of the seeds and a schematic body plan
of all species complete the account. The most notable change is all former subspecies of D.
stolonifera are elevated to species rank. i.e. D. stolonifera subsp. humilis , D. stolonifera subsp.
monticola, D. stolonifera subsp. porrecta, D. stolonifera subsp. prostrata , and D. stolonifera
subsp. rupicola are treated as the new species D. humilis, D. monticola, D. porrecta, D. prostra¬
ta, and D. rupicola. Drosera stolonifera subsp. stolonifera is retained as a more narrowly defined
species, Drosera stolonifera. Drosera purpurascens (that has formerly been attributed to D.
stolonifera subsp. stolonifera ) is now found identical with what has been described as D.
stolonifera subsp. compacta, so D. purpurascens is the correct name of the latter taxon if regard¬
ed a species different from D. stolonifera. (JS)
Sirova, D., Adamec, L., and Vrba. J. 2003. Enzymatic activities in traps of four aquatic species
of the carnivorous genus Utricularia. New Phytologist. 159: 669-675.
By examining the enzymatic activity in four species of aquatic Utricularia (U. vulgaris, U.
australis, U. foliosa, U. aurea), the authors were able to detect the presence of acid phosphatase
inside the bladders. It is unclear if the creation of this enzyme is due to the plant itself, or organ¬
isms living inside the bladders and resisting consumption. (BR)
Wistuba, A., Carow, T., Harbarth , R, and Nerz, J. 2005, Heliamphora pulchella, eine neue mit
Heliamphora minor (Sarraceniaceae) verwandte Art aus der Chimanta Region in Venezuela. Das
Taublatt. 53/3: 42-50.
The new species Heliamphora pulchella is described. Previously regarded as a form of
Heliamphora minor , this new species is distinguished by the long (5 mm, 0.2 inches) hairs on
the inside of the pitcher. (BR)
58
Carnivorous Plant Newsletter
Writings from the Readership
Unexpected Hybrids of Spectacular Bladderworts Species
Miloslav Studnicka • Botanic Gardens Liberec • Purkynova 1 • 460 01 Liberec • Czech
Republic • botangarden@volny.cz
Keywords: cultivation: hybridization, Utricularia.
Introduction
The genus Utricularia is divided into 35 sections. Two of these sections — Iperua P. Taylor
and Orchidioides DC. — have some of what are arguably the most spectacularly large and beau¬
tiful species in the genus. For examples, the beautiful white- flowered U. alpina and the red-flow¬
ered U. quelchii are both from section Orchidioides', the huge peltately-leaved U. nelumbifolia
and hugely-flowered U. humboldtii are from section Iperua. Since these various species do not
usually grow together in nature, and presumably have different pollinators, hybridization is not
known from the wild.
It is nearly as difficult for attempts at hybridization to be made in cultivation. It is unfortu¬
nately rare for bladderworts from these sections to flower in cultivation. Synchronous flowering
of different species from these sections to flower at the same facility is altogether sporadic.
However, over the past fifteen years we have had a few opportunities to attempt hybridizations
of these plants at the Botanic Gardens of Liberec.
To date, the hybrids are all infertile, but they expand vegetatively. The first successful hybrid
cross was made between U. humboldtii (female) and U. quelchii (male) in 1990. The resulting
plant is very similar to U. humboldtii, but it is distinctly smaller, leaves being mostly about 16
cm high. Following the nature of the seed parent, it does not produce bulbs but does produce aer¬
ial offshoots. The first flower of the hybrid, produced in August 1997, had 43 mm broad lower
lip (see Figure 1 ). This hybrid makes an effort to produce flowers only sporadically, and that is
why it is a poor horticultural plant (and not worth, in my opinion, being registered as a cultivar).
Nevertheless, the successful hybridization of the species belonging to different sections of their
genus is very interesting (see Discussion section, below).
The hybridization between Utricularia quelchii (female) and U. praetermissa is quite dif¬
ferent, in being an easier plant to grow and flower. Accordingly, I have established a cultivar
name for it ( Utricularia ‘Jitka’; Studnicka (2005)). I have also tried to make the same cross, but
inverting it so that U. praetermissa was used as the female plant. The results were plants that
produced flowers that look very much like the archetypical U. praetermissa ! I suspected auto¬
gamy to be occurring (which would be very unusual in the section Orchidioides) and confirmed
this later, by means of an isolated specimen of this species which produced viable seed.
Yet another hybrid was produced in 2000 between U. alpina (female) and U. campbelliana
(male). This hybrid is characterized by flowers 54-59 mm across, and which are pallidly cerise
(see Figure 2). The lower lip of the corolla is saddle-shaped, because it droops down on both
sides. There is a golden-yellow palate splotch on the lower lip. The upper lip is longer than the
upper calyx lobe. Flowers hang pendently on very thin, 20 cm tall peduncles. There is a whorl
of three bracts on the stem, quite like in U. campbelliana. The plant is tuberous, and much small¬
er than U. alpina — spathulate leaves are at most only 7 cm long and 1 cm broad. Cultivation of
this hybrid is unfortunately difficult, as in U. campbelliana. That is why the novelty, which
bloomed for the first time in 2002, is probably unsuitable to most growers, and is not deserving
of cultivar status.
Volume 35 June 2006
59
Figure 1 : A cross made between U. humboldtii (female) and U. quelchii.
Figure 2: A cross made between U. alpina (female) and U. campbelliana.
60
Carnivorous Plant Newsletter
Discussion
Of the two very similar species U. alpina and U. praetennissa , autogamy was only observed in
U. praetennissa.
It is striking that the cross U. humboldtii x U. quelchii produced viable seed, since the two par¬
ent plants are in different sections in the genus Utricularia. From this we could deduce that the sec¬
tions Iperua and Orchidioides are closely related. Alternatively, we could deduce that U. humboldtii
is more properly identified as a species from the section Orchidioides. Furthermore, if U. humboldtii
were transferred to section Orchidioides, the section Iperua would have to be given a new name
since it will have lost its type species. Should we accept this transferral, we find that the section
Orchidioides (including U. humboldtii) and the section that was previously called Iperua occur in
geographically separated areas. Section Orchidioides would occupy the Andes, Venezuelan moun¬
tains, and the northern part of the Brazilian Shield. The section that was once called Iperua (but with¬
out U. humboldtii would be given a new name) would be restricted to the southern part of the
Brazilian Shield. The primaeval Shield was divided into the northern and southern parts by the much
younger Amazonian Basin. This geo-historical separation could be the reason for the segregation of
the sections Orchidioides and Iperua. We find various analogies in both sections, like tuberous
species (U. geminiloba in IperuaV) and species symbiotic to Bromeliaceae , with aerial shoots.
The counter-hypothesis, i.e. transferring U. quelchii into section Iperua is not as compelling
since the geographic separation of the two new sections is not as clean. Furthermore, there are a
number of other species in section Orchidioides (such as LJ. endresii) that are clearly closely related
to U. alpina.
Perhaps in these ruminations, we should also take into consideration botanical intuition. If you
compare the strange solid spathulate dark-green leaves of both U. humboldtii and U. quelchii, they
seem to be native sisters; and the hybridization discussed here seems to be less surprising.
References:
Studnicka, M. 2005, New cultivar: Utricularia ‘Jitka’. Carniv. PI. Newslett. 34: 27-28.
Volume 35 June 2006
61
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Phone: 704-399-3045 Cell: 704-458-8538 v
http://personal.atl. bellsouth.net/clt/f/l/flytrapl /index, html
Contact: DAVID CRUMP Price list - S.A.S.E.
Cook's Carnivorous Plants
P O Box 2594 Eugene Oregon 97402
541/688-9426
http //www flytraps com/
Specializing in the strange and unusual plants since 1992
Ovet 150 varieties available foi purchase at all times, as well as live moss and your needed supplies
By appointment on site sales . Shipping yeai round worlrto/ide!’
Give us a call from 9Am to GPm Pacific Coast time Monday tlmi Friday check us out on the wel> anytime
Email support through cooksqflvtiaps.com
Carnivorous Plant Nursery
Promoting environmental stewardship through
engaging educational materials and low-cost, high-quality carnivorous plants.
• Carnivorous Plants
• Bog Orchids, Bog Plants
• Activity Book for Carnivorous Plants
• Bugs in the Bog Board Game
• Free Monthly Drawing
• OnLine Shopping and Mail Order
16128 Deer Lake Road, Derwood, M D 20855
http://vvww.ca rnivorousplantnursery.com
Phone: 301-519-7505 Email: info(o)carnivorouspIantnursery.com
62
Carnivorous Plant Newsletter
1
Cascade Carnivores
Carnivorous Plant Nursery
• Drosophyllum lusitanicum
- Yes, we ship Dewy Pine plants - live delivery guarantee
• Nepenthes
- highland species are our specialty
• Drosera, Pinguicula, Utricularia,
Byblis, Heliamphora, VFTs and more
Visit us on the web - http://www.cascadecarnivores.com
email - support@cascadecarnivores.com
The New York Botanical Garden Press
Drosera (Droseraceae)
Mireya D. Correa A.
and Tinia Regina dos Santos Silva
Flora Neotropica Monograph 96
In Spanish with an English abstract
0-89327-463-1
72 pages
$25.00
Send all orders to:
The New York Botanical Garden Press
20odi Street and KazimirufF Boulevard
Bronx. New York 10458-5126 USA
718. 817. 8721 • tax 718. 817. 8842
nybgpress@nybg.org • www.nybg.nrg
When in Northern California Visit
California
Carnivores
Specializing in insect-eating ana other exotic plants
* Open all year, call ahead in winter
* Commercially cultivated
* Over 500 varieties on display
* On site & domestic mail order
* We ship potted plants
2833 Old Cravemlein Hwy South, Sebastopol, CA 95472
(707)824-0433 Price List-SASE
Visit os on the Internet at http://californiacsrnivores.com
fRIFFI® PARK
257 PERRY ROAD, KEYSBOROUGH, VIC, 3173, AUSTRALIA
PHONE: 61 (0)3 9769 1663, I lam to 5pm Monday to Thursday ONLY
FAX: 61 (0)3 9769 1663, 8am to 8pm, 7 days per week
EMAIL: triffids@triffidpark.com.au WEB: www.triffidpark.com.au
Owned and operated by Colin and Tina Clayton. Managed by Donna ClaytonSmith.
FOR ALL YOUR CARNIVOROUS PLANT REQUIREMENTS INCLUDING:
PLANTS, POTS, LABELS, BOOKS, SEEDS, SPHAGNUM MOSS AND PEAT MOSS
WRITE, PHONE, FAX OR EMAIL TRIFFID PARK FOR A FREE COLOR
MAIL ORDER CATALOGUE OR VIEW OUR COMPREHENSIVE WEB SITE.
You are most welcome to visit Triffid Park but please organise this with us first,
as inspection is by appointment ONLY.
Volume 35 June 2006
63
§,*m
Vy r , J.';
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11