NOV 1 1 1983

JNJW YORK

(»4^a

CARNIVOROUS
PLANT
NEWSLETTER

Official Journal of the
International Carnivorous
Plant Society

Volume 12, Number 3
September 1 983

COVER PHOTOS

Front: Rosette plant and lower pitcher of Nepenthes sanguined.

Back: Lower pitchers of Nepenthes sanguined.

See story, page 65. Photos by Roger Shivas

The co-editors of CPN would like everyone to pay particular attention to the
following policies regarding your dues to the ICPS.

All correspondence regarding dues, address changes and missing issues should be
sent to Mrs. Pat Hansen, 3321 Hamell Rd., Fullerton, CA 92635. DO NOT SEND TO
THE CO-EDITORS. Checks for subscriptions and reprints should be made payable to
CSUF FOUNDATION-1 CPS.

All material for publication, comments and general correspondence about your
plants, field trips or special noteworthy events relating to CP should be directed to one
of the co-editors. We are interested in all news related to carnivorous plants and rely
on the membership to supply us with this information so that we can share it with
others.

Views expressed in this publication are those of the authors, not necessarily the editorial staff
Copy deadline for the December issue is October 1 5, 1 983.

CO-EDITORS:

D. E. Schnell, Rt. 1 , Box 1 450, Pulaski, VA 24301

J. A. Mazrimas, 329 Helen Way, Livermore, CA 94550

T. L. Mellichamp, Dept, of Biology, UNCO, Charlotte, NO 28223

Leo Song, Dept, of Biology, California State University, Fullerton, CA 92634

Seed Bank: Patrick Dwyer, St. Michael’s Episcopal Church, 49 Killean Park, Albany,

NY 12205, USA.

BUSIN ESS MANAGER: Mrs. Pat Hansen, c/o The Fullerton Arboretum

PUBLISHER: The International Carnivorous Plant Society by the Fullerton Arboretum,
California State University, Fullerton, CA 92634. Published quarterly
with one volume annually. Printer: Kandid Litho, 129 Agostino Rd., San
Gabriel, CA 91776. Circulation: 656 (160 new, 496 renewal). Dues:
$10.00 annually, $15.00 foreign. Reprints available by volume only.
®1983 Carnivorous Plant Newsletter. All rights reserved.

58

Carnivorous Plant Newsletter

News and Views

TOM CARPENTER (21334 Park Mount
Drive, Katy, Texas 7 7450) writes: Would
you expect a natural CP population in
the middle of the South’s largest city? A
small population of Drosera brevifolia is
maintaining itself in an open, regularly
mowed sandy area adjacent to a Houston
city park. The waiter has visited the loca¬
tion several times over the past three years
during which time the population seems
to have neither enlarged nor declined.

Recently, the writer contacted Mr. Di-
anni in East Texas (CPN News and Views,
Dec. ’82) concerning a Sarracenia alata
population about to be destroyed. Mr.
Dianni still intends to bulldoze, drain,
and convert the location to grazable
pasture. However, as of March ’83, the
population was still intact. The writer
found the plants to be heavily infested
with Exyra moths. Also present were Dro¬
sera capillans and Utncularia subulata.

From CHRIS LUHDORFF (6473 Sierra
Drive, Coulterville, CA 95311): Have
you ever been stumped on what to give
as a gift to a friend or loved one? In my
family, a home-made gift has more senti¬
mental value than something from the
store, and so I suggest the following: If
you live in the mountains, as I do, go out
and collect pieces of “bark-like” wood
that can be whittled. Carve out a cup¬
shaped indentation, then fill it with soil.
This makes a good planter for Drosera.
If the indentation is deep enough, you
can plant Dionaea or Cephalotus in it.

If you live near the beach, you can col¬
lect shells that are deep enough to fill

with soil and use as planters. Use your
imagination. See what you can come up
with! (Ed. note: Please respect any regula¬
tions regarding collection of materials from
protected areas.)

JOE MAZRIMAS reports: I had over 60
orders for the new K. Rondo book and all
orders were processed on time. It is not
economically feasible for me to process
single orders, so for all those who would
like to order a book, please send an inter¬
national Money Order of $10.00 to:

Dr. Katsuhiko Rondo
Faculty of Integrated Arts & Sciences
Hiroshima University
Hiroshima 730, JAPAN
For those of you who like to send pic¬
tures and slides for publication in CPN,
we urge you to send us pictures of top
quality. It costs us the same price to pub¬
lish the good ones as it does the bad qual¬
ity pictures. You may send color or B & W
photos,* 35 mm slides or larger size
slides. It is better for these pictures to be
slightly overexposed (about 14 stop) than
to be dark since some darkening occurs
in the reproduction process. Most of the
features in these images should be in
sharp focus and plants should fill nearly
the entire field for best details to show. It’s
difficult to see the plants if the picture is
taken many feet away. All pictures will be
returned at the request of the author.
Please include a couple of sentences
about the plants in the picture, especially
where the plants were found and the
ecology of the area.

Pollination of Sarracenia purpurea 60

Preservation of Nepenthes pitchers 62

Nepenthes miribilis variation 64

Nepenthes of Gunung Ulu Rali 65

Sarracenia hybrids 68

Gland movement of Drosera capensis 69

Glossy photos, please.

Sarracenia purpurea

drawing by Ron Fleming

In this issue:

Volume 12 • September 1983

59

International Carnivorous Plant Society

SEED BANK*

Patrick Dwver (St. Michael's Episcopal Church,
49 Killean Park, Albany, NY 12205)

Byblis liniflora (8)
Darhngtonia califomica
Dwnaea muscipula
Drosera aliciae (4)

D. auriculata (10)

D. burkeana (10)

D. capensis

D capensis (narrow) (5)
D capillaris
D. indica (10)

D. intermedia
D. lovellae (10)

D. montana (2)

D. peltata
D. pygmaea (4)
l) rotundifolia

D. spathulata (Formosa) (8)
D. spath. (Kansai)

D. spath. (Kanto)

D. spath. (white fl.)
Drosophyllum lusitanicum (10)
Nepenthes gracilis
N. khasiana
N. mirabilis
Pinguicula alpina (10)

P. Corsica (10)

P. grandiflora palhda (6)
P. vulgaris
P. vulgaris bicolor (2)
Sarracenia alata ( 1 2)

S. flava
S. leucophylla
S. purpurea purpurea
S. rubra (3)

S. X chelsonii (8)

S. flava X minor (2)

S. leuco X alata (5)

S. rubra X leuco (5)

*For details on how to send or order seed, please see CPN, March 1983, page 4.

A Preliminary Report on the Pollination of
A Sarracenia purpurea in a Forest-Swale Ecotone

by

Wendv O'Neil, Director, Natural Area Registry Program,

The Nature Conservancy, 531 N. Clippert, Lansing, MI 48912

edited by Lam' Mellichamp

Abstract

This studv was conducted during the
summer of 1979 while the author was a
student at the University of Michigan Bio¬
logical Station, near Pellston.

Sarracenia purpurea flowers (161 flowers)
were studied for a period of 29 hours dur¬
ing anthesis over a nine-day interim. Vis¬
itors to the flowers were collected in hope
of establishing the pollinator(s) of the
plant. Members of the Flalictidae (a cos¬
mopolitan family of small black or brightly
metallic solitarv bees) primarily were
caught and thought to be actting as pol¬
linators. Bombus terricola (bumblebee) fe¬
males were also collected and may play a
minor role in pollination. Other insects,
including flies and beedes, were found as
visitors, although they were unlikely pol¬
linators.

Introduction

Sarracenia purpurea L. is an insectivorous
plant that grows commonly in Northern
sphagnum bogs. Although first described
in 1601, it was not until a century' later that
Dr. Michael Sarrasin of Quebec sent speci¬
mens of this plant to Tournefort in Paris
(Harper, 1918). Work on Sarracenia has fo¬
cused on the insectivorous nature of the
plant and the ecological interactions with¬
in the pitcher-shaped leaves (Istock et al.,
1975).

Pollination studies, by comparison, have
been fewer and less detailed. Schnell (1978)
mentions a small species of Bombus as a
possible pollinating agent for Sarracenia
rubra. In 1965, Adrienne Mandossian of
Michigan State University completed a
doctoral thesis on the life history of Sarra¬
cenia purpurea in a sphagnum bog habitat

60

Carnivorous Plant Newsletter

in Northern Michigan. In it, she concluded
that major visitors to her plants were: Sar-
cophaga sarraceniae (a fly), Bombus impatiens,
B. grisecollis, B. terricola, B. vagans, Apis mel-
lifera and several species of ants. Results of
her study also indicated that cross-polli¬
nation provided a larger amount of seed
set than self-pollination, and that non¬
pollination provided almost no seed set.

In Sarracenia purpurea, llowers are borne
on peduncles 3-5 mm high with five dark
red petals that droop. There are live per¬
sistent sepals and three bracts. The flower
goes through three main stages: young,
medium, and old. Young flowers are those
that have recently opened and still bear
petals and anthers. Medium-aged flowers
have dropped petals and most of the
stamens. Older flowers have no petals or
stamens left. At functional maturity the
flower is held upside down at the tip of the
bent-over flowering stem. During anthesis,
nectar is secreted at the base of the barrel¬
shaped ovarv. Should nectar be unremoved
from the plant it collects and runs down the
ovary.

Materials and Methods

The study site was a wet forest-swale (low
dunes) ecotone along the shore of Lake
Michigan located at Grass Bay, in section
25 of T38N, RIW, and section 30 ofT38N,
RIW, Cheboygan County, Michigan.

Field observations were made by check¬
ing flowers along a 290 meter transect that
ranged from 5 to 10 meters in width.
Flowers were observed between 0830 and
1630 hours. 161 Sarracenia flowers were
checked for visitors during 29 hours of
observation spread over 9 days.

Flower visitors were normally not visible
until the petals were cautiously pulled back.
If an insect was inside, the petals were re¬
placed and a net placed over the flower.
The time was recorded and the insect cap¬
tured upon exiting from the flower. It was
necessary to allow the insect to leave the
plant of its own accord so that pollen pres¬
ent on its body would be the result of its
own activity rather than a by-product of
the capture procedure. Captured insects

were placed in a potassium cyanide kill¬
ing vial and later pinned for a voucher col¬
lection.

Insects were examined for pollen loads
under a dissecting microscope. Pollen loads
were removed from some of the halictid
bees and treated with glycerine jelly, 9596
alcohol and methyl green using a method
described by Rapp (1969). The loads were
then examined under a compound mi¬
croscope to determine whether or not
Sarracenia pollen was present.

Flowers that were at the young, medium
and old stage of anthesis were bagged with
cotton gauze and left covered from mid¬
afternoon July 6, 1979 until the morning
of July 8, 1979. Nectar was sampled from
these llowers with micropipets and placed
on a pocket refractometer in order to
measure sugar concentration.

To establish that llowers were actually
being pollinated, a return visit to the site
was made on August 12th to check for
capsule and seed production.

Results

A total of sixteen types of insects were
found visiting the inner portion of the
Sarracenia flower. The stage of anthesis
seemed to affect the nature of visitor.
Twelve insect types were found visiting
the young and medium-aged plants where¬
as four types were found on old flowers.

Bees gathered from the flowers were
determined as follows: one visitor was an
Osmia sp. (Megachilidae); two bumble¬
bees were identified as Bombus terricola
(Apidae) females; and 43 Halictidae, al¬
though not officially identified, were
thought to be the same species of small
bee.

One of the Bombus was found inside a
flower. It had forced its way in between a
petal and the stigma. The other Bombus
was observed crawling in over the stigma
and exiting between a petal and sepal.
Microscopic examination revealed white
Sarracenia pollen on both bees. Only on
two other occasions were Bombus seen
near the plants. Of these, one was in flight
and the other approached a pitcher flow-

Volume 12 • September 1983

61

Preservation of Nepenthes Pitchers by
Freeze Drying

Roger G. Shivas, Botanv Department,

National University of Malaysia, Bangi, Selangor, Malaysia

Dried and pressed herbarium speci¬
mens of Nepenthes do not clearly exhibit
many of the characteristics of the living
plant. In particular dried pitchers often
lose their original shape and form. How¬
ever, it is often the shape of the pitcher
which has been used to identify the many
species and natural hybrids of Nepenthes.

Curators of zoological museums have
preserved small animals bv the method
of freeze drying. This method of preser¬
vation has a wider application in micro¬
biology and packaging certain foodstuffs.

The method of freeze drying requires
leaving the material to be preserved in a
vacuum at a temperature of about -50° C.
The period of time that material is left in
the freeze drier depends upon the nature

of the material. Bacterial cultures are
generally freeze dried for 4 hours, whereas
a small bird may require a week of freeze
drying.

Nepenthes pitchers were freeze dried for
24 hours using the apparatus available at
the Zoology Department, National Uni¬
versity of Malaysia. The pitchers main¬
tained their original shape and to a certain
extent their colour. Pitchers preserved by
this method were obtained from plants
of Nepenthes macfarlanei, Nepenthes san¬
guined, Nepenthes albo-marginata and Nepen¬
thes ampullaria. The freeze dried pitchers
were only slightlv brittle and durability
might be obtained by coating the pitchers
with a clear varnish paint or sprav.

Pollination (from page 61)
er and at the last second avoided it.

The Halictid bees were found inside the
pitcher blossoms repeatedly. Several times
they were found perched amongst the stam¬
ens, with their heads facing the base of the
ovary. Most commonly, they were found
wandering around inside on the stvlar disc
which was covered with pollen. Usually
they exited a pitcher plant blossom be¬
tween the petals and the sepals, or between
the stylar disc and petal, nearlv always mak¬
ing a short flight to the sepals and resting
there while combing gathered pollen onto
the tibia of their hind legs. On two occa¬
sions, bees were observed leaving the flow¬
er across the stigmadc surface.

The other bee collected was not seen
on any other occasion. It was an Osmia sp.
and carrying pollen on the underside of
the abdomen.

Twenty-eight members of the family
Sarcophigidae, probably Sarcophaga , were
seen on pitcher blossoms. These flies were
rarely found in the young flowers. Instead
they were noted on flowers of medium and

old ages.

Other insects collected from the flowers
included fireflies (Lampyridae) - old flow¬
er; click beetle (Elateridae) — medium flow¬
er; wasps (Sphecidae) — young flower;
midges; ants; cranefly; mosquito. None of
these was considered to be involved with
pollination.

When Halictid pollen loads were ex¬
amined, Sarracenia pollen was determined to
be white in color. Its stephanolcolporate
pollen grains were easily identifiable
under high-power magnification. Four
out of fourteen bees examined had pol¬
len from at least one other species in addi-
| tion to Sarracenia pollen.

Nectar results showed a sugar concen¬
tration ranging from 17.2% (wt/wt) in old
flowers to 35.2% (wt/wt) in young flowers.
The nectar production was also found to
decrease with flower age.

Discussion

Since the Sarracenia flowers were already
opened when the study period began it
was not possible to undertake manipula-
(Continued on page 74.)

62

Carnivorous Plant Newsletter

Plate A Lower pitcher of Nepenthes macfarlanei prior to freeze drying.

Photos by Roger Shivas

Plate B Freeze dried pitchers ot Nepenthes macfarlanei.

Volume 12 • September 1983

63

Nepenthes mirabilis variation

by Bruce Lee Bednar (25 Lake Court Loop, Ocala, FL 32672)

The mirabilis complex, this common
form of Nepenthes, is variable. The dif¬
ferent forms often have been given separate
identities as species, but eventually were
recognized as variations of a single specie.
Some of the plants in circulation have
acquired incorrect names accidentally, or
sometimes fraudulentlv to increase trading
value to put as much mileage on them as
possible. The problem of old synonyms
occurs with plants mostly out of Europe
where many mirabilis are still labelled phyl-
lamphora , an abandoned name not used in
decades.

The plant in CP trade labeled kampo-
tiana is not at all the original plant de¬
scribed by Lecomte. The true species is not
believed to be in cultivation at this date.
Instead, the bogus “kampot” is thought to
be a natural hybrid between mirabilis and
thorelii , and has appeared on newer collec¬
tors’ lists unofficiallv labelled x lecouflei. To
add more wood to the fire, there is doubt to
the authenticity of “thorelii-long green.”
Leconrte’s paper on thorelii is difficult to
obtain, if it’s still around at all. Many feel
“thorelii-long green" is the Thailand mir¬
abilis, and the pubescent leaved, squat pit-
chered “short round” is the true species.
Il this is so, many hybrids using thorelii as
a parent are mislabeled and plants like x
hachiyo and x effulgent koto would be inter¬
grades, not hybrids.

I’ve seen very little information on the
species anamensis described bv Macfarlane
as a highland variety. The one in cultiva¬
tion seems also to be a mirabilis variant.
The light green pitcher, maroon and green
wavy leaves are too similar to the other
mirabilis types of the Indo-China area.

Two other plants have been brought to
my attention as being bogus. One which
seems to have been renamed in Australia
somewhere along the line is geoffrayi, anoth¬
er plant of Lecomte. I have not seen the
bogus tvpe which is typical mirabilis, ob¬
viously labelled so as to increase its pop¬
ularity. The true specie geoffrayi is not at
all similar to mirabilis, and there would
be no room for confusion in identifying
the two.

Smillesii is another example, truly a
mirabilis with a “new name.” The plant has
been around for vears; rename it and once
again the plant sells.

A new monograph is needed for the
genus Nepenthes. The mirabilis complex
needs much work itself. Until a standard is
achieved, many Nepenthes collectors will
remain lost. In the next few years many
plants not in cultivation will become avail¬
able. It is very important to be able to
properly identify them so others can con-
fidendy rest assured what they have la¬
belled is really what they have. Any com¬
ments?

NEW PLANT AND
SEED SOURCES

Catalogue Price

Stock

Catalog Price

Stock

Carnivorous Supplies
P. O. Box 179

Albion Park

N.S.W. 2527

AUSTRALIA

International
Reply Coupon

CP

Seed

The Cottage Nursery
Moortown Road,
Nettleton

Lines LN7 6HX

ENGLAND

P

CP

64

Carnivorous Plant Newsletter

Nepenthes of Gunung Ulu Kali

Roger G. Sliivas

Botany Department, National University of Malaysia, Bangi, Selangor, Malaysia

The Genting Highlands are situated
about 40 km northeast of Kuala Lumpur in
tropical Malaysia. The highest mountain
in this part of the range is Gunung Ulu
Kali (1800 nt). Presently the area is being
developed as a tourist resort and boasts
four hotels, a goll course, and a casino.

Gunung Ulu Kali is olten shrouded in
cloud and rainfall in the area is high.
The temperature ranges between the ex¬
tremes of hot days (30° C) and cool
nights (10° C).

The vegetation on the ridges and crests
is upper montane moss forest character¬
ized by stunted trees of Dacrydium and
Leptospermum, several species of Rhodo¬
dendron and other ericaceous shrubs and
many epiphytic orchids. The top soil is
peat, which is often covered in dense mats
of sphagnum moss.

Three species of Nepenthes have been
recorded from Gunung Ulu Kali, Nepen¬
thes macfarlanei. Nepenthes gracillima var
major and Nepenthes sanguinea (Stone,
1981). During 1982 and 1983, I made
several trips to Gunung Ulu Kali to ob¬
serve and collect Nepenthes.

Nepenthes macfarlanei is a robust plant
with waxy leaves and was the most abun¬
dant plant species in the moss forest.
This species was readily distinguished by
its cream-coloured, crimson-spotted
upper pitchers and large, ovate (10 to 15
cm high and 5 to 8 cm wride), mottled red
lower pitchers. Plants often reached
lengths greater than 4 m.

Nepenthes gracillima , although not as
plentiful as Nepenthes macfarlanei, was
easilv identified by its dark purplish-
black, cylindrical pitchers (10 to 20 cm
high and 1 to 3 cm wide). Mature plants
were vines greater than 5 m in length. The
form of this species found on Gunung
Ulu Kali most closelv fits Nepenthes gracil¬
lima var major (Ridlev, 1924).

Nepenthes sanguinea was rarely found on
the ridges and crests of Gunung Ulu Kali.

However, this species was found along
roadside embankments at an altitude 300
m lower than the summit. Nepenthes
sanguinea was conspicuous with its wide
peristome (1 to 2 cm) and cylindrical
pitchers (10 to 20 cm high and 3 to 6 cm
wide). The colour of the pitchers varied
from jiale red and green to a dark, uni¬
form blood red.

Literature Cited

Ridley, H.N. (1924) The Flora of the Malay
Peninsula, III London, L. Reeve & Co.,

p. 22.

Stone, B.C. (1981) The summit flora of
Gunung Ulu Kali (Pahang, Malaysia).
Federation Museums Journal, Kuala Lum¬
pur, Volume 26 (Part I), N.S.

(See front and back covers as well as
photos, pages 66-7).

Volume 12 • September 1983

65

Upper pitcher of Nepenthes macfarlanei.

Photo by Roger Shwas

Lower pitcher of Nepenthes macfarlanei.

Photo by Roger Shivas

Carnivorous Plant Newsletter

Figure 1 — The smaller plants are green
S. alata X minor (North Carolina), and (he
larger plants are the red-throated S. alata
X S. minor (Ok.efenok.ee).

Figure 2 — S. alata red-throated X S. f lava
heavy-veined.

Photos above by Steve Clemesha.

Distinctive purplish-black pitchers of Nepenthes gracillima.

Photo by Roger Shivas

67

Volume 12 • September 1983

THE INFLUENCE OF DIFFERENT
FORMS OF SARRACENIA SPECIES
ON THEIR HYBRIDS

bv Steve Clemesha

(Lot 6 Skinner Close, Avocado Heights, Woolgoolga, NSW Australia 2456)

Two Sarracenia hvbrids that I made
about 14 vears ago were A alata X A
minor and A alata X A flava. In both cases
the common green form of A alata was
used while the S. minor was also the com¬
mon form — a plant from North Carolina,
and the A flava was the heaw-veined
form.

The two hvbrids matured. The A alata
X S. flava are tall green plants. The pitchers
are slender and thev have some red veins.
The pitcher tops and hoods show the
influence of S. flava but thev are not as
large or spectacular as in that species.
The plants produce good pitchers right
through the growing season and like the
A alata parent, plants produce offsets
freely.

The S. alata X A minor does not grow
very large. Most pitchers are 20 cm or less
tall, rather like A minor in shape but lack¬
ing its colour. Onlv two or three small
light window spots are found on the upper
pitcher back, and these could be easily
missed unless looked for. The main con¬
tributions A alata made in this hybrid
were its green colour and freely offsetting
habit. In all the two hvbrids are easily
grown, but are not very colourful.

Four vears ago, 1 repeated the crosses
using the form of S. alata with red inside
the hood, the large Okefenokee Swamp
form of A minor and the form of A flava
that has a red pitcher with a green hood
that is red veined. The plant of S. flava is
from Florida.

The hvbrids using these parents are
strikingly different from those made
earlier. The A. alata X A. flava is now nearly
mature. Its pitcher shape is much like that
of the earlier hybrid but the young pitch¬
ers bv the time they are open are red from
the pitcher rim to the base. The hood at
first is green with red veins inside and out.

After a few days, the green becomes gold¬
en and soon after this the hood becomes
solid red inside and out. The colour is
particularly intense on the inside of the
hood where it is almost black-red. Late
summer and autumn pitchers colour as
fully as spring ones. In winter the pitcher
tops die but the live bases remain red.

In the A alata X A minor hvbrid the
plants are also more colourful. The upper
third of the pitchers and hoods are red¬
dish-brown, and the hood interiors are
solid red, though this is not easily seen
because of the A. minor- like shape. The
upper pitcher back has numerous white
spots.

The most outstanding feature of this
hy'brid, however, is its size. Pitchers are
50-55 cm tall. They taper from their
narrowest point at the base to their widest
at the pitcher mouth where they are 5 cm
from front to back of live pitchers. The
hood also is large. The pitchers of these
plants are taller and broader at the top
than either parent.

I have made others using the Okefeno¬
kee S. minor. So far onlv the S. X areolata
cross with it is near maturity. It differs
little from the S. alata cros but ntav not be
as tall.

I have crossed the red-mouthed A alata
with A leucophylla and with A purpurea ssp.
venosa. The A leucophylla cross differs little
from other plants of A X areolata I have.
The pitcher tops are a bit darker and the
hood interior is red as in the A alata.
Because the A leucophylla colour is not im¬
proved, this hybrid is not very' attractive.

The cross between the red-mouthed A
alata and A purpurea ssp. venosa shows the
influence of the colour of A alata in the
colouring of the hy'brid hood interior.
The colour is soon masked as the whole
pitcher reddens but the hood interior

68

Carnivorous Plant Newsletter

remains darker than in hybrids where the
green form of S. alata was used.

Having grown the Okefenokee Swamp
S. minor, the red- mouthed S. alata and
red form of S. alata (with the green lid
which is red-veined) for more than ten
vears I have known for some time that
their distinctive characteristics are not the
result of growing conditions. They come

true from seed it the plants are selfed
and it is now evident that their distinctive
characteristics are passed on to at least
some of their hybrids and in some cases
with vcrv striking results.

In all cases the plants are grown out¬
side in lull sun. Winters here are very
mild with rare frosts.

(See color photos, page 67.)

A Photographic Study of the
Rapid Movements of Stalked Glands of the
Cape Sundew Drosera capensis

Glenn Elzinga, John D. Beebe, and Randall Van Dragt
Biology Department, Calvin College, Grand Rapids, Michigan 49506

The movement of stalked glands and
leaves of sundews is well known. The
first record of modes of behavior of
stalked glands and leaf blades in the genus
Drosera can be traced to Dr. Roth in 1782
(Lloyd, 1942). Lloyd documented slow
leaf blade movement of D. capensis in a
sequence of frames from a time-lapse
motion picture. Bopp and Weber (1981)
have published photographs of slow leaf
blade bending of D. capensis in a hormone
regulation study. Williams (1976)
reported on rapid stalked gland move¬
ment and slow leaf bending movement
in photographs of D. intermedia taken bv
James Kowalchuk. In our present studv
we present photographs of D. capensis
(Figure 1) which show the rapid stalked
gland reponse (Figure 2) to the presence
of an ether anesthetized fruit fly ( Droso¬
phila melarwgaster).

An anesthetized fruit 11 v was placed on
the lateral extreme of the outer discal
stalked glands of a D. capensis leaf blade
(Figure 2a). The terminology is that of
Llovd (1942) reporting on the work of
Behr who divided the stalked glands of
D. rotundifolia into three groups pro¬
gressing from the 1. marginal glands,
2. the outer discal glands, to the 3. discal
or central group of glands. In one hour
(Figure 2b) the fly had been coated with
mucilage and had been carried medially

a short distance bv the extreme outer
discal glands. At two hours (Figure 2c) the
fly has been carried to the lateral margin
ol the discal or central glands. The ex¬
treme bending of the outer discal glands
became apparent and the bending of the
marginal glands can be seen. During the
next two hours (Figures 2e and f) the
position ol the fix' did not change appreci¬
ably; however, the progressive bending
of the marginal glands continued. In a
period of five hours the fruit fly was car¬
ried from a lateral position to a central
position on the leaf blade and the outer
discal and marginal glands have shown
extreme bending.

The plants were two year old cuttings
in 7.5 cm plastic pots containing a peat-
vermiculite mix. The plants were grown
in a glass case with a loose fitting cover
and two 20 watt cool white fluorescent
lights 5 cm from the top of the plant and
10 cm from the top of the pot. The plants
were watered from the base with 1 to 2
cm of deionized water and allowed to drv
slightly before the next watering. Between
photo taking sessions the plants were
covered with a glass beaker in order to
maintain high humidity.

The photographs were made using a
Canon FI camera and Kodak Panatomic
X film. The D. capensis habit photograph
(Figure 1) was made using a 25 mm exten-

Volume 12 • September 1 983

69

sion tube and a 50 mm macrolens with a
lens to subject distance of 2 cm. The plant
was 9 cm from a masked Sunpack ring
Hash. The gland movement series (Figs.
2-7) involved the use of a bellows ex¬
tended 105 mm with a 50 mm macrolens
at a distance of 2 cm from the lens to
the subject and two flashes — one direct
30 cm from the subject and one reflected
from a white card 20 cm from the sub¬
ject. All exposures were fl 6 and 1/60 sec.
Negatives were printed on Kodak Poly¬
contrast Rapid II RC F paper.

Literature Cited

Bopp, M. and f Weber. 1981. Hormonal
regulation of the leaf blade movement
of Drosera capensis. Physiol. Plant. 53:
491-496.

Lloyd, F. E. 1982. The carnivorous plants.
Chronica Botanica Company, Wal¬
tham, Mass.

Williams, S. E. 1976. Comparative sens¬
ory physiology of the Droseraceae —
the evolution of a plant sensorv svstem.
Proc. Amer. Phil. Soc. 120 (3): 187-204.

Figure 1. A two-year old cutting of Drosera capensis grown under lights in an artificial soil mix.

70

Carnivorous Plant Newsletter

Figs. 2-7. Stalked gland movement of Drosera capensis in response to an anesthetized fruit
fly during a five hour period.

Volume 12 • September 1 983

71

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Carnivorous Plant Newsletter

Volume 12 • September 1983

73

Pollination (from page 62)

tions of plants which would have yielded
information about cross-pollination, self-
pollination, and actual seed set. Thus it
will be necessary to attempt interpretations
ofthe results while bearing in mind that
the project was started a bit late for yield¬
ing conclusive results.

In Mandossian’s thesis (1965), she re¬
ports four species of Bornbus including
Bornbus terricola as frequent visitors to Sarra-
cenia flowers. She observed them to be ex¬
tremely active through most of the day,
moving constantly from flower to flower,
never staving long inside. She also report¬
ed entrace to be over the stigma. Man-
dossian observed large pollen loads on
the Bornbus.

The two Bornbus terrkola females caught
in this study were the only two found in
the llowers during the entire study period.
They did have a fair amount of pollen
grains on their bodies. Of the bee visitors,
Bornbus comprised only 496 of total insect
visits. While it is possible that Bornbus are
pollinating Sarracenia at this site, it seems
likely that thev play only a very minor role,
if at all.

Halictid bees seem to be the primary
visitors to Sarracenia llowers.* They ac¬
count for 9596 of flower visitations. While
they were observed to occasionally exit
flowers across the stigma, and thus it could
be suggested that the pollen which they
gather while crawling about inside the
llowers is deposited on the stigmatic sur-
lace and effects self-pollination, it is more
reasonable to assume that since they
normally enter the llower across the stig¬
matic surface (and exit between petal and
stigma) that they are cross-pollinating the
flower by depositing pollen as they enter.

Mandossian (1965) found that only 2996
of flowers produced seeds that had insect
visitors excluded and were left untouched.
Sixty-four percent of the flowers which she
self-pollinated produced seed, while 9796
of the flowers that were cross-pollinated
produced seed. This indicates that Sarra-
cema purpurea is self-compatible and can
produce seed even in the absence of pol¬
linators. It also shows, however, that insect

visits that result in self-pollination can be
beneficial. Unfortunately, it is impossible
to evaluate the relative extent to which the
Halictids cross- and self-pollinate Sarrace¬
nia in the present study. Interestingly,
Mandossian (1965) does not mention
Halictids as visitors at all. It is possible
that this difference could be accounted
for bv habitat differences.

Decidedly, this study should be repeated
for more conclusive results. It would be
necessary to start much earlier — as early
as the emergence of buds in order to
determine quantitatively what is occurring
in the flowers. This would enable bagging
of plants, exclusion of visitors, outcross¬
ing and self-crossing experiments. Specific
plots could be designed wherein records
of the visits to each plant could be kept.
The period of stigma receptivity and pol¬
len maturity should be determined as well.
It might not be a bad idea to check the
flowers a few more times at night - to elim¬
inate or establish the possibility of moth
visitors. Despite Mandossian’s thesis on
the life history of Sarracenia (1965), it might
be a productive study if Sarracenia flowers
were watched at a bog site and at this
swale site during the same season. Thus,
it could be established whether or not vis¬
itors to Sarracenia are specific to the plant
and differ by latitude or whether they are
specific to the habitat in which the plants
exist.

Literature Cited

Harper, Roland M. 1918. The American
pitcher plants. J. Mitchell Soc., Sept.,
110-125.

Istock, C.A., S.E. Wasserman, and H. Zim
mer. 1975. Ecology and evolution of
the pitcher-plant mosquito: I. Popula¬
tion dynamics and laboratory response
to food and population density. Evolu¬
tion 29: 296-312.

Kapp, R.O. 1969. How to know pollen and
spores. Win. C. Brown Co. Publishers,
Dubuque. 249 pp.

Mandossian, A.J. 1965. Some aspects of
the ecological life history of Sarracenia
purpurea. Unpubl. PhD. thesis, Michi¬
gan State Univ.

Schnell, D.E. 1 978. Systematic flower stud¬
ies of Sarracenia. Castanea 43: 21 1-220.

74

Carnivorous Plant Newsletter

Review of Recent Literature

Earley, L. S. 1983. The alluring pitcher
plant. (Illustrations bv Carol Lerner).
Wildlife in North Carolina 47 (6):2-3.

This hall page popular article briefly
describes the four sarracenias in North
Carolina, but also features a one and a
half page spread of Carol Lerner’s
excellent line drawings. (DES)

Earley, L. S. 1983. Two days in John
Green’s Swamp. Wildlife in North
Carolina 47 (6): 14-21.

This is an excellent “color piece” de¬
scribing exploring the Green Swamp
in modern times, as well as an inter¬
esting historical review of the times
and trials of the Swamp. The article is
accompanied by good color photos,
including a typical savanna and a clump
of Sarracenia flava. (DES)

Grollman, J. Flesh Eaters at South Ken¬
sington. New Scientist, April 28, 1983.

The author describes the displays of
CP at the Natural History Museum in
London, which were exhibited be¬
tween April 21 and June 3 in sealed
glass cases. The Carnivorous Plant
Society is sponsoring this exhibition
to familiarize visitors with these fas¬
cinating plants.

Kondo, K. and Y. Yaguchi. 1983. Stomatal
responses to prey capture and trap
narrowing in Venus’s llvtrap (Dionaea
muscipula Ellis). II. Effects of various
chemical substances on stomatal re¬
sponses and trap closure. Phyton 43:
1-8.

Application of various plant growth
substances either stimulated or inhib¬
ited trap closure, and some caused the
trap to remain closed for two or more
days. Apparent lluid tension changes
in subepidermal tissues suggest that
water movement and transpiration rate
are involved in trap movement. (DES)

Lounibos, L. P., C. Van Dover, and G. F.
O’Meara. Fecundity, autogeny, and
larval environment of the pitcher-

plant mosquito, Wyeomyia smithii.
Oecologia (Bed.) 55 (2): 160-164,

1982.

The authors collected pupae and 4th
instar larvae from Alabama Sarracenia
purpurea pitcher plants. They found
that the females which emerged varied
greatly in the production of mature
eggs and didn’t have any relationship
to the density of larvae found in in¬
dividual pitchers.

WANT ADS

When submitting Want Ads, please be
sure to print clearly for best results and
to eliminate mistakes. Please circle the
correct letter before each item (Want,
Trade, Sell or Buy). Want ads are limited
to carnivorous plants, terrariums, green
houses and moss. There is a charge of
ten cents per item, with no limit to the
number of items you may submit per
issue.

Send coin or check to:

Arboretum, Want Ads
California State University
Fullerton, CA 92634

Richard Chinnock (3316 Old Dirkwood
Dr., Virginia Beach, VA 23452). To sell:
Sarracenia plants, too many to list. Pingui-
cula lutea plants, Dionaea muscipula plants.
Send SASE for list and prices.

Curtis Yax (12 Division, Apt. 1, Oneonta,
New York 13820). Trade: Gemmae of
pygmy sundews. Send SASE.

C.G. Antoni (330 West 22 Street, New
York, N.Y. 10011). Trade: Cuttings of
N. X coccinea with any Nepenthes cuttings
except N. coccinea, khasiana, and kampotiana.

Note: All individuals or organizations selling,
trading or buying CP are advised to be cognizant
of certain restrictions under the U.S. ESA and
international CITES for certain species (see edi¬
torial).

Volume 12 • September 1983

75