VOLUME 1 1 , Number 2

JUNE 1982

CARNIVORC

PLANT

NEWSLETTE

International Carnivorous
Plant Society

Official Journal of the

Volume 1 1 , Number 2
June, 1 982

COVER

Pinguicula macroceras. Photo taken by Jurg Steiger in March, 1979.

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, addresschangesand missing issuesshould be
sent to Mrs. Kathy Fine, c/o The Fullerton Arboretum, Dept, of Biology, California State
University, Fullerton, CA 92634. DO NOT SEND TO THE CO-EDITORS. Checks for
subscriptions and reprints should be made payable to CSUF FOUNDATION-
ARBORETUM.

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 Sept, issue is Aug. 1 , 1 982.

CO-EDITORS:

D. E. Schnell, Rt. 4, Box 275B, Statesville, NC 28677

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

T. L. Mellichamp, Dept, of Biology, UNCC, Charlotte, NC 28223

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

BUSINESS MANAGER: Mrs. Kathy Fine, 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
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®1982 Carnivorous Plant Newsletter. All rights reserved.

AUG 0 9 1982

30

Carnivorous Plant Newsletter

_ JOKK

^.iTAMICAI nARDFN

News and Views

ALLEN LOWRIE (6 Glenn Place, Dun-
craig 6023, Western Australia) has just
returned from Borneo and reports that
his Nepenthes cuttings arrived safely using
the following method: The cuttings were
placed in plastic hags with only enough
water to mist the sides of the bags after
they had been sealed 20 minutes.

BILL NETHERBY (P.O. Box 1, Mt.
Herman, CA 95041) recently persuaded
the garden editor of the Santa Cruz Sen¬
tinel to do a spread on CP. The articles
by Keith Muraoka appeared in two in¬
stallments and generally described the
habits of CP that we know so well. We
thank Bill for calling attention to CP and to
CPN where interested plant growers can
expand their interests.

TONY REA announces that the San
Francisco Flower Show will take place
August 27-29 at the Hall of Flowers in
Golden Gate Park. Bay Area CPN mem¬
bers will be sent an application. See
CPN 10(4), 101 (1981) for details.

Our intrepid correspondent DALE
SPEIRS of Calgary, Alberta sent an inter¬

esting article from an 1875 issue of Bo¬
tanical Gazette (1:66-67). It is actually
a review of other current periodicals in
which is mentioned an article in the Oc¬
tober American Naturalist by Prof. W. J.
Beal entitled Carnivorous Plants. The usual
genera are listed along with Silene spp.
which often crops up in the older litera¬
ture in this respect. However, another
genus we had not before considered was
also discussed in some detail, this being
the species Martynia proboscidea. As often
happens over time, the genus has now
been changed to Proboscidea in the family
Martyniaceae, two rather common and
widespread species being P. louisiamca
and P. parviflora in North America. The
genus is most recognized for its pecul¬
iar fruit from which the common names
“unicorn plant” and “devil’s claws” are
derived. The fruits are often pickled as
food. Prof. Beal apparently felt that the
densely glandular character of vegetative
portions of the plants (similar to Silene in
this respect) was an indication that it was
carnivorous, and he went so far as to
(Continued on page 4b.)

SEED BANK*

Patrick Dwyer (St. Michael’s Episcopal Church,

49 Killean Park, Albany, NY 12205)

$.75 per packet

Byblis liniflora (2), Cephalotus folhcularis ( 1 ), Darlingtonia californica ( 3),Dionaea muscipula, D. aliciae (9), D.
auriculata (8), D. arcturi (3), D. binata (1), D. binata multipda (2), D. brevifoha (2), D. burkeana (3), D.
burmannii(S), D. capensis, D. capensis (narrow), D. capillaris (2), D. filiformis filiformis (3), D. indica (3), D.
intermedia, D. linearis ( 1 ), D. macrophylla (2), D Montana ( 1 3), D. natalensis ( 1 ), D. peltata (6), D. planchonii

(3) , D. pygmaea (1), D. rotundifolia, D. rotundifolia (Oregon) (2), D. spathulata (7), D. spath. (Kansai), D.
spath. (white fl.) (2), D. stenopetala (1), D whittakeri (3), Nepenthes ampullaria (6), N. gracilis (10), TV.
khasiana, N.mirabilis (13), TV. rafflesiana (12), Pinguicula primuliflora (1), P. vulgaris, Sarracenia flava, S.
leucophylla, S. minor ( 1 4), S. purpurea heterophylla (6), S. purpurea purpurea, S. purpurea venosa ( 1 0), S. rubra

(4) , S. flava X oreo (1), S. leuco Xalata (1), S. psitt. X minor (4), S. sp. mix (3), Utricularia racemosa (1), U.
subulata (1).

‘For instructions on how to send or order seed, see CPN March 1982, page 4,

Volume 11 • June 1982

31

Continued from March 1982

From

SINNESORGANE IM PFLANZENREICH

bv Gottlieb Haberlandt

Insectivores: Dionaea muscipula

Translated by Carla R. Powell
Departments of Chemistry and Foreign Language
Lebanon Valley College, Annville, PA 17003

Upon treatment with zinc chloride iodine
solution3, these underlying cell wall layers
exhibit only a light yellowish- grey coloration,
while the epidermal outer walls of the leaf
blades up to the cuticle immediately turn an
even shade of blue. There can scarcely be a
doubt that, just as in the case of Aldrovanda,
the differing chemical behavioral of the our
er walls of the hinge cells correlates with
their flexibility and elasdcity.

The radial longitudinal walls (side walls) of
the hinge cells are considerably thickened.
This is easily observed on crosswise micro¬
tome sections ( Plate VII, Fig 2) and tangen
dal longitudinal secuons through the hinge
(Fig. 5). On tangential longitudinal sections
one also sees that the upper and lower per¬
ipheral portions of the longitudinal walls
are thinner and, as a result the cell lumina
are enlarged at these points The longitudi¬
nal walls turn blue and swell greatly when
treated with zinc chloride iodine’. Even with¬
out further treatment after the protoplasts
have been fixed with chroma osmium acetic
acid8, a delicate cross- striadon of the some
what swollen longitudinal walls is observed
resulting from the presence of more numer¬
ous plasmodesmata between the adjoining
hinge cells (Plate VII, Fig 3)9. The sloping
lateral walls, which border on die hinge cells
above and below, are just as thin as the
radial walls of the adjacent epidermal cells,
corresponding to them.

The inner walls of the hinge cells are
again quite thick (Plate VII, Fig. 1); but the
thickening does not usually extend all the
way to the upper and lower transverse edge
of the inner wall. These walls also are trav¬
ersed by plasmodesmata , such that the pro¬
toplasts of the hinge or sensory cells are con¬
nected not only with one another, but also
with the protoplasts of the central cell bun¬
dle

This cell bundle, surrounded by 15-16
hinge cells in the form of a ring, consists of
longitudinally stretched cells with perpendi¬
cular, or only slightly slanted transverse walls
(Plate Vf Fig 10). The bundle i$ as a rule
two cell layers high, and on median length
wise sections, three cell layers wide In the
case of an exceptionally weakly constructed
brisde, I found a single longitudinally stretched
cell in place of the enure bundle. The con¬
tent of these cells includes a living plasmatic
body with an ellipsoidal nucleus. The na
ture of their walls is peculiar. They are
more or less thickened and with relatively
weak magnification, appear as though they
are covered with rounded pits (This cor
rects the mistaken description of these cell
walls in the first edition of this book p
113.) Renewed investigation, however ex¬
plained this appearance Many strongly re
fractive granules, or sinuate platelets are
embedded in the middle lamellae in a sin¬
gle layer. To judge by their reactions they
consist of a cutose substance10. On thin mi¬
crotome sections (5-10 fi m thick one sees
very clearly, with sufficiendy strong magnifi¬
cation, that the rows of granules are covered
on both sides by smooth cellulose lamellae,
which correspond to the layers of secondary
thickening (Plate VII, Figs 1, 2). Such cutin
granules also occur in the partition walls be¬
tween the hinge cells and the central cells
It is worth noting that, except for the Ion
gitudinal walls only the transverse walls in
side the hinge cells exhibit this construc¬
tion. The upper- and lowermost transverse
walls, through which the cell bundle bor¬
ders on the adjacent tissue consist of ref
atively pure cellulose and have no granules.
On the whole, those characteristic wall prop¬
erties are not so much determined by the
cell boundaries, as much more by the up¬
per and lower boundaries of the hinge Here
the height of the hinge is equal to the
height (or length) of the inner walls of the

32

Carnivorous Plant Newsletter

epidermal hinge cells. The walls of the cells
existing within these boundaries and the cell
parts exhibit the qualities just described.
Therefore; it may be concluded that these
characterisuc wall properties are connected
with the function of the hinge. The impor¬
tance of the deposidon of numerous cutose
granules in the cellulose wall is a complete
mystery.

In the first edition of this book, I con¬
sidered the central cell bundle in question
to be the mechanical tissue of the entire
hinge, and compared it, as far as its func¬
tion is concerned, with the central body of
leaf pulvini. This conception was based on
the incorrect assumption that the radial lon¬
gitudinal walls of the hinge cells were thin
and delicate, such that the presence of a
central mechanical cell bundle would be con¬
ceivable Now, however; the laminar hinge
cells in general appear to be so thick- walled
that a special mechanical cell bundle in the
middle of the hinge probably is superfluous.
Since there • are plasmodesmata between the
hinge cells and the central cell bundle; the
most important function of the bundle must
be to transmit to the pedestal, or lamina
the state of excitation caused by deforma
tion of the hinge cells".

Fig. b. A large leaf of Dionaea in its spring form.
An arrow points to one of the bristles (trigger
hairs) on the surface of the leaf. This partic¬
ular leaf has eight bristles as many larger leaves
do. Leaves with six bristles would have one
bristle, in place of the centrally located pairs
of bristles, located on each side of the blade
exactly between the pairs of bristles nearest
the midvein. Haberlandt’s Fig. 10 in Plate VI
is a longitudinal section through the base of
one of these bristles.

4. The undermost part of the tactile bris¬
tle is formed from a cylindrical pedestal
which is approximately as high as it is wide.
It consists of thin- walled plasma rich paren
chymatous cells, which are occasionally faint
ly collenchymatous and are surrounded by
an epidermis having rather elongated cells.
It is worth noting that only the innermost
layers of the moderately thickened outer walls
of these epidermal cells are stained blue by
zinc chloride-iodine, while the outer layers
take on the same yellowish- gray color as the
outer walls of the hinge cells, without be
ing cuticularized This indicates that when
the brisde is more severely bent, both the
actual hinge and the pedestal are slightly
bent In agreement with this the pedestal
is not broadened at its base. Indeed it is
inclined to be thinner here than it is fur¬
ther above (See Haberlandt 1896, Fig 207;
Goebel s illustration is incorrect in this is¬
sue) By more severe bending of a brisde
under a microscope, one can readily observe
that, in fact, the pedestal is also bent ( Hab
erlandt, 1896, Fig 208). And indeed this
bending is provided for in the structure of
the pedestal When the brisde is severelv
bent as it is particularly after the leaf halves
close; its function evidendy is to prevent a
too great possibly permanendy damaging
deformation of the hinge Howevet with
weaker impulses this is probably the onlv
place bending will occur. The strongest bend
ing will undoubtedly occur in the hinge
zone characterized bv the ring- shaped con¬
striction

I must now discuss briefly the nature of
the deformations of the protoplasts of the
hinge cells caused by their bending.

On observation of a median longitudinal
section through the bristle, one sees that the
hinge furrow is actually none other than a
crease in the thickened outer walls ( Plate VI
Fig 10 and Plate If Fig 1). Each hinge
cell or sensory cell has in its very thick
outer wall a much thinner cross- striation
When the hinge is bent the convex side is
gready stretched while the concave side
forms a fold at this point ( Haberlandt 1896,
p 208). When the convex side is stretched
the portion of the plasma membrane adjoiir
ing the thin wall striation is subjected to ex¬
treme stretching and this deformation of
the protoplast, or plasma membrane is evi¬
dendy greater here than at any other place
on the cell deformed bv the bending. On
the concave side the nature of the mech-

Volume 11 • June 1982

33

Fig. c. Plate IV fig. 10 from the second edition
of Sinnsorgane. Plate VI is the same in both the
first and second edition of the book except for
this drawing (c.f. Plate VI Fig. 10). Note that
Haberlandt changed his drawing in the second
edition to indicate that all walls of the cells in
the middle laver (k) are suberized instead of
just the uppermost walls. Because of the much
lower quality of printing of the plates in the
second edition photographs of Plate VI are
taken from the first edition.

anical demands placed on the plasma mem¬
brane cannot be so precisely defined In the
case of a weaker bending, it may have been
subjected to a longitudinal pressure On the
other hand, with a strong bending where
by the thin portion of the outer wall folds
inward, a longitudinal stretching of the plas¬
ma membrane combined with radial com¬
pression probably occurs In any case how¬
ever; the important difference in the thick¬
ness of the outer walls of the hinge cells
serves to localize the deformation on a very
specific part of the plasma membrane That
the stretched plasma membrane is especial¬

ly sensitive at these places; that is under
the only slightly thickened hinge striations
of the outer walls, is an obvious surmise1'.

It is possible to explain more clearly the
workings of the hinge furrow when the bris¬
tle is bent, if we calculate and compare with
one another the bending moments of two
hollow cylinders which we imagine to repre¬
sent the outer walls of the hinge cells at
their thickest and thinnest points. The for¬
mula for the bending moment1* of a hol¬
low cylinder in which the contents are no
glected is: F(r^ + r*)/4. In this formula F
represents the cross- sectional area and r and
r, the radii of the hollow cylinder differing
by the thickness of the wall ( Schwendenei;
1874, p 23). I determined the following val¬
ues for F rp and r2 on the basis of meas¬
urements of a drawing of the median longi¬
tudinal section through the hingq carried
out by means of a [camera lucidaj. In the
case of a hollow cylinder whose wall is
formed from the thickest portion of the
outer walls of the hinge cells, F = 405 mm2,
r = 23 mm and r, = 20 mm The size of
the bending moment is 93960, accordingly.
In the case of a hollow cylinder whose wall
thickness is the same as the thin portion
of the outer walls of the hinge cells F =
122 mnf, ^ = 20 mm and i; = 19 mm
The bending moment is 23180. The actual
hinge cell is therefore considerably more
weakly constructed than the thickened por¬
tions of the wall adjoining on both sides
Simple observation of the dimensional rela
tionships themselves makes this obvious In¬
deed the ratio of the bending moments is
4:1, i e, the thickened portions of the walls
of the hinge cells are approximately four
times less flexible than the thin ones

If the relationship between the structure
and function of the tactile bristle presented
above is correct, then it should make no
difference to the resulting motor response
whether the stimulation, namely the bend¬
ing of the bristle is caused by a solid ob
ject, or bv a fluid striking against it, for
example a jet of water Contradicting this
Darwin (1876, pp 263, 264) says that “ drops
of water or a thin, intermittent stream fall
ing down from a certain height onto the
filaments” produces no stimulation Of course
it was previously indicated bv Goebel '1891,
p 202)., that raindrops usually run off the
thin brisdes without moving them That will
certainly be the case if the direction in which
the drops are falling coincides with the lon-

34

Carnivorous Plant Newsletter

Fig. d. upper portion — a longitudinal section
through the sensory cells of the trigger hair
(c.f. Plate VI, Fig. 10). Lower right — a pro¬
jection of the thickest part of the outer cuti¬
cle and cell wall illustrating one-fourth of the
hollow cylinder used to compute the moment
of inertia (1 ) of the thickest point. Lower left

— a projection of the thinnest part of the out¬
er cuticle and wall illustrating one-fourth of
the hollow cylinder used to compute the mo¬
ment of inertia (I.) of the thinnest point. 1 =
Ffr2 + r2 )/4 where F = the cross-sectional area
of the projected cylinder, r = the outside ra¬
dius and r = the inside radius.

gitudinal axis of the upright bristle; or forms
an acute angle with it If howevet; a lat
eral jet of water is aimed at the bristles
then, as Balfour has shown, the motor re
sponse occurs immediately".

The tactile brisdes of Dionaea must, accord¬
ing to the preceding material, be considered
as very completely developed percepdon or¬
gans. If on the whole they are the most
complete and most highly specialized organs
of this species, then this presumes a very
long period of evoludon in the course of
the phytogeny. This seems all the more like

ly since no other primitive brisdes occur on
die leaf which might have been the phylo¬
genetic predecessor of the tactile bristles' .

With regard to sensors’ physiology, it is
highly interesting that sense perception is
not exclusively limited to those highly spe-
cialized organs developed for that purpose
Various researchers, such as Darwin, Munk
Goebel and others agree that the upper side
of the leaf blade is also sensitive to mech¬
anical stimulation, albeit to a much lesser
extent. Movement may also be caused by a
wound

Volume 11 • June 1982

35

I\.ClClClICC&

A. Batalin 1877. Mechanik der Bewegung der
insectenfressenden Pflanzen Flora 35: 54-
58; 105-111; 129-154.

MA Curtis 1834. Enumeration of the plants
around Wilmington, N. C Boston Journ
Nat Hist 1: pp 123-127.

G Darwin 1876. Insektenfressende Pflanzen
trans V Carus ( Stuttgart). p 250.

Sydenham Edwards 1804. Dionaea Musapula.
Venues Fly-Trap Curtis Botanical Maga
zine 20, plate 785.

J. Ellis 1770. A botanical description of the
Dionaea musapula or Venus’ s flytrap a
newly discovered sensitive plant. Nova Ac¬
ta Regae Societatis Scientarum Upsaliensis
1: 98-101.

K. Goebel. 1891. Pflanzenbiologische Schil -
derrungen (Marburg), pp. 57-72.

G Haberlandt 1896. Phvsiologische Pflanzen-
anatomie IL (Auf}, pp 481 482.

F. Kurtz 1876. Zur Anatomie des Blattes der
Dionaea muscipula. in Reichter and DuBois-
Reymond Arch Anat und Physiol ( Leip

zig, pp 1-29.

J. Lindley. 1848. An Introduction to Botany,
4th ed (London).

J.M. Macfairlane. 1892. Contributions to the
history of Dionaea muscipula Ellis. Contri¬
butions of the Bot Lah of the Univ. ol
Pennsylvania 1 7-44.

F.J. F. Meven 1839. Neues System der Pflan-
zenphysiologie. 3: 545.

H. Munk 1876. Die electrischen und Be
wegungsterscheinungen am Blatte der Dionaea
muscipula. in Reichter and DuBois- Reymond
Arch, fur Anat und PhvsioL ( Leipzig pp
30-203.

T. Nuttal 1818. The Genera of North Amer¬
ican Plants ( Philadelphia), p 277.

C. A Oudmans 1859. Over de prikkelbaar-
heid der bladen van Dionaea musapula El¬
lis. Verslagen en Mededeelingen der k.
Akademie van Wetenschappen 9: 320.

S. Schwendener. 1874. Das mechanische Prim
zip im anatomishen Bau der Monokoty-
len ( Leipzig, p. 23.

ENDNOTES

by Stephen E. Williams, Department of Biology
Lebanon Valley College, Annville, PA 17003

1. This very important experiment clearly
shows that the “endpiece” or lever portion
of the trigger hair which lies above the sen¬
sory cells of the “hinge” is not a part of
the sensory system. Benolken and Jacobson
(1970, J. Gen. Physiol. 56: 64-82), although
they were unaware of Munk’s experiments,
did the same type of experiment — only in
reverse. They sliced away the podium bit by
bit and found that the electrical signals could
be produced until they cut into the cell lev¬
el of the hinge. Each of these experiments
indicates that some or all of the cells at the
hinge are sensory in nature, as Haberlandt
states as the major thesis of this part of his
work. However, it is also possible that other
cells distal to the hinge cells have a sensory
function as well, as has been pointed out by
M. Williams and Mozingo (1971, Amer. J.
Bot. 58: 538).

2. Haberlandt apparently is clouding the is¬
sue here. There is little doubt that the sen¬
sitive response is usually triggered by the
hairs and that Munk and Darwin believed
that to be so. The question is really whether
the ability to respond is also in other parts
of the leaf which, although sensitive, are

more difficult to stimulate because of their
structure or other factors or whether the
hair is the only sensitive part of the leaf.
They found other parts of the plant to be
sensitive, a fact which is of some physiolog¬
ical interest and which Haberlandt recog¬
nizes in later parts of this chapter.

3. Zinc Chloride-iodine solution was a test
for various polysaccarides which could be
identified by the color they develop when
treated with this solution.

4. Walls in this middle layer, which Hab¬
erlandt states are suberized, stain with Sudan
black B in the same way that the walls of
the endothermal layer in the Drosera tenta¬
cle stain (S. Williams, 1976, Proc. Amer. Phil.
Soc. 120: 187-204). Lloyd (1942, Carnivorous
Plants, p. 188) stated that he could “find very
little if any suberization” in this layer. The
methods used by both Lloyd and Haberlandt
to detect suberization are not given. The Su¬
dan stain, which dissolves in hydrophobic
lipid structures such as cuticular wax, indi¬
cates that these walls are impermeable to
water and are like the Casparian strips found
in the walls of many glandular structures.
There are important changes in the drawing

36

Carnivorous Plant Newsletter

of this structure that were made in the sec¬
ond edition of Sinnsorganne (c.f. Fig. 10 and
Fig. c).

5. The great significance of these structures
is precisely due to the fact that they do not
have a function. Like the leg bones of the
python or our appendix they may be rem¬
nants of structures that had functions in an¬
cestral organisms. Nearly all plant glands
have an endodermis with Casparian strips of
some sort. The presence of such a structure
in the trigger hair of Dionaea is important
evidence that this purely sensory structure
may be derived from a secretory structure
very much like the Drosera tentacle. The ho¬
mology of the Drosera tentacle with the Dio¬
naea and Aldrovanda trigger hairs is discussed
in detail elsewhere (S. Williams, 1976, Proc.
Am. Phil. Soc. 120: 187-204).

6. The correctness of Haberlandt's state¬
ment is easily confirmed by looking at the
electron micrograph of the 1-2 pm thick cu¬
ticle of this region in Fig. 12 of M. Williams
and Mozingo (1971, Amer. J. Bot. 58: 232-
539).

7. Again M. Williams and Mozingo’s (1971,
Amer. J. Bot. 58: 232-539) observations agree
with those of Haberlandt rather than those
whose he is critical of. There are clearly no
pores. However, the denticulations he reports
are reported to be cuticular vesicles by M.
Williams and Mozingo, who were observing
sections of tissue less than 90 nm (0.09jL/m)
thick in contrast to Haberlandt’s 4fj m thick
sections. It is quite possible that denticula¬
tions which were attached to the cuticle have
been cut through in a way that makes them
look like vesicles in thin sections. The fact
that they remain attached to the cuticle when
Haberlandt hydrolized away the cellulose
walls with sulfuric acid supports this inter¬
pretation (Plate VI, Fig. 11). Electron micro¬
scopic studies are best done in parallel with
light microscopic work.

8. This mixture is also called Fleming’s
fixative. It has several variations containing
different concentrations of chromic acid, os-
mic acid and acetic acid in water (Huma-
son, 1967, Animal Tissue Techniques, p. 19).

9. These plasmodesmata were hvpothesized
to exist by Benolken and Jacobson (1970, J.
Gen. Physiol. 56: 64-82) to explain the phy¬
siological data they observed. Haberlandt’s
drawings (Plate VI, Fig. 10 and Plate VII, Figs.
2, 3) clearly support their hypothesis but M.
Williams and Mozingo (Amer. J. bot. 58: 538)
report that these plasmodesmata do not ex¬

ist. They do report large numbers of plas¬
modesmata connecting the sensory cells -
an observation which Haberlandt failed to
make. Because of the grat care with which
he did his work Haberlandt’s observations
should not be taken lightly. However, M.
Williams and Mozingo’s (1971, Fig. 8) lon¬
gitudinal sections show some plasmodesma¬
ta in thinner portions of the wall but not
the large number reported bv Haberlandt.
It would be interesting to know just what
Haberlandt was looking at.

10. I was unable to find Sudan black B
positive walls in this part of the hair, even
though it was in this region that I was ex¬
pecting them. I started looking because of
the analogy of these cells and the hinge cells
with the sensory cells of Aldrovanda. If these
granules are cutin they should take the Su¬
dan stain. This observation should be re¬
peated.

11. A thorough investigation of the distri¬
bution of plasmodesmata throughout the trig¬
ger hair and adjacent leaf blade is in order.
This is the best way that the pathway over
which action potentials spread can be de¬
termined. The pathway Haberlandt suggests
here is questionable on the basis of a com¬
parison of his description of the distribution
of plasmodesmata with that of M. Williams
and Mozingo (1971, Amer. J. Bot. 58: 532-
539).

12. Haberlandt’s emphasis on the cell mem¬
brane sounds very up to date. It is quite
likely that the membrane is the site of re¬
ception rather than a number of other or¬
ganelles mentioned in far more recent stud¬
ies of the trigger hair.

13. The “bending moment” refers to the
moment of inertia of the cross section of the
tubular outer wall and cuticle of the hair
(see Fig. d). The moment of inertia (I) to¬
gether with a constant called Young’s mod¬
ulus (E) determines how much force it takes
to bend the hair. One can determine a “stiff¬
ness factor” (S), where S = El, that deter¬
mines how difficult it is to bend any partic¬
ular cross section through the hair. (See
Thomas, Calculus and Analytic Geometry, Read¬
ing, Mass., pp. 550-551). Haberlandt is com¬
paring the relative ease of bending the hair
at two points along its length — that is, at
the narrowest point of the hinge (point b)
and at a nearby point (point a or c) where
the wall and cuticle of the hair are very
thick. A way to compare these is to find the
ratio of the stiffness factors (S^VSj,) at the two

Volume 11 • June 1982

37

points. Since StySb = EcIc/Et)Ih it follows that
= 1^/^ if the Young’s modulus (E) is
the same at both points on the hair. The
Young’s modulus (E) depends on how easy
the material in the wall and cuticle is to
stretch. Haberlandt has assumed it is equal¬
ly easy to stretch per unit cross-sectional
area at both points and so he has computed
I,/Ib = 4. His conclusion is that it is four times
as easy to bend the hair at the narrow place
than at the broad places so that bending will
tend to take place there. Haberlandt may have
used a slide rule since his calculations are not
exact, as may now quickly be ascertained with
a calculator. The calculated values for I have
far too many significant figures for the data
anyway, so the precision is more than enough
to justify the approximate 4: 1 ratio of stiffness

factors.

14. If the tendency of the trigger hair to
require more stimuli for closure as the stim¬
uli are delivered at less frequent intervals is
added to this discussion, this would be a
very accurate account of this phenomenon
(c.f.s. Williams, 1980, CPN 9: 65, 75).

15. For a discussion of the evolution of
this structure see S. Williams, Proc. Am. Phil.
Soc. 120: 187-204.

Acknowledgments: Thanks are due to Dr. S.
L. Jacobson and Dr. Y. Heslop-Harrison for
reviewing the manuscript and to Mrs. Carla
Lange for bibliographic help. Dr. James
Scott of the Lebanon Y'allev College Foreign
Language Department made major contribu¬
tions to the translation.

Plate VII, Left Side

Dionaea muscipnla
(Microtome sections)

Fig. 1. Part of a radial longitudinal secdon
through the hinge of the tactile bris¬
tle; the sensory cell is pictured togeth¬
er with the protoplast (x 700).

Fig. 2. Cross section through the hinge of the
tactile bristle; inside the ring of radial
sensory cells is the central bundle, with
cutin particles embedded in its cell
walls. Fixed with alcohol, stained with
paracarmine (x 660).

Fig. 3. Partition wall between two sensory cells;

cross sectional view. The wall is pen¬
etrated by plasmodesmata; fixed with
chrome-osmium acetic acid, stained
with paracarmine (x 880).

Fig. 4. Part of a cross section through the
hinge of a tactile bristle; the sensory
cells touch at the upper ends, such that
on the outside, the cross sections
through the dowrn-tumed parts of the
epidermal cells bordering above are
still visible (X 660).

Fig. 5. Tangential longitudinal section (saggi-
tal section) through the sensory cells
of the tactile bristle (x 700).

Fig. 6. Cross section through the upper part
of the stimulator of the tactile bris¬
tle (x 480).

Plate VI, Right Side

Dionaea Muscipula

Fig. 10. Longitudinal section through the bas¬
al portion of the tactile bristle (x 440).

Fig. 1 1 . Surface view of the hinge of a tactile
bristle. The cuticle of the stimulus-
sensitive cells is very finely denticu¬
lated on its inner side.

Drosophyllum lusitamcum

Fig. 14. The same, [surface view of two cells
of the epidermal glandular layer] of
a sessile gland.

Fig. 16. The same; [isolated protoplasts of the
lateral glandular cells of a parietal ten¬
tacle, viewed from the side] the cells
in question were situated more toward
the apex (x approx. 900).

Fig. 17 Isolated proplasts of an apical glandu¬
lar cell (X approx. 900).

Fig. 18 Surface view of some lateral glandular
cells (X approx. 900).

Drosera longifolia

Fig. 19. Isolated protoplasts of two lateral glan¬
dular cells. After treatment with dilute
sulfuric acid and staining with toluidine
blue (X approx. 1000).

Drosera dichotoma

Fig. 21. Surface view of tw'o lateral glandular
cells of a cluster. Treated with Javelle
water.

Fig. 22. Surface view of some apical glandu¬
lar cells.

38

Carnivorous Plant Newsletter

LEFT SIDE

PLATE VII

Volume 11 • June 1982

39

PLATE VI

RIGHT SIDE

40

Carnivorous Plant Newsletter

A PHOTOGRAPHIC PRIMER OF VARIANTS
OF SARRACENIA RUBRA WALT.

bv Donald E. Schnel!

As was the case in the previous two
editions of this primer series, text re¬
marks must be kept brief, an especially
difficult task in this very controversial
species. Most regular CPN readers and
students of Sarracenias are well aware of
the problems of taxonomy of Sarracenia
rubra. A selected bibliography is append¬
ed and readers are encouraged to read
as many of these papers as available for
details and still further important refer¬
ences. (Reprints of the author’s papers
cited are still available in limited quan¬
tities.) Throughout this discussion, the
author’s nomenclature will be used; it
is becoming more widely and generally
accepted (primarily references 5 and 8
with additional documentation in 6 and

7)-

However, the other systems of no¬
menclature must also be studied and
considered, and all papers present im¬
portant concepts and insights. Wherrv (9)
first declared one of the S. rubras a
separate species ( S . jonesii), but was later
content with subspecies status (10). Bell
(1) thought the jonesii plants were but
a variety and somewhat extended the
putative range; he recognized no other
infraspecies. McDaniel (4) felt that there
were no significant taxonomic differ¬
ences at all. The Cases (2, 3) however
preferred to split S. rubra into three
separate species and one subspecies.
Range maps can be found in references
1, 2, 3, 4 and 5, each drawn with the
author’s taxonomic intentions in mind.
The maps in 3, 4 and 5 are quite similar.

The numbers preceding the paragraphs
below correspond to the figure numbers.

1) Flower of S. rubra ssp. rubra, quite
representative of all the flowers in dif¬
ferent subspecies, the only significant
difference being in size in some popu¬
lations. Note the red petals (may be pale
red to maroon), reflexed sepals in older
flowers. The fragrance of all sspp. is uni¬

formly sweet or pleasant. The undersides
of the petals in most plants of anv ssp.
is tan to tan-green in color and often has
a linear red streak characteristic of the
species.

2) S. rubra ssp. rubra, eastern North
Carolina. The range of this ssp. is the
eastern Carolinas extending inland to the
sandhill counties. Within the species as
a whole, there is considerable seasonal
plcomorphism of developing pitchers,
early spring pitchers tending to be small
to etiolated with many curved forms
having prominent ala. This example is
quite small although flowering size, has
earlv spring pitchers and grows in a less
than ideal habitat that dries during the
summer. Plants in such areas remain
more juvenile.

3) S. rubra ssp. rubra , Lexington Co., SC.
This example of the same ssp. as in Fig. 2
is growing in a prime habitat: an open,
sunnv area in sands' soil and in a large
seep that is constantly wet. Note the very
robust pitchers that approach other sspp.
in Fig. 4 and 5 in character. One of the
problems in studying the species is the
wide variation due to local factors, vari¬
ations that tend to be neutralized in
transplant and cultivation experiments.
Also, the differences between sspp. in
S. rubra are not at all sharply discon¬
tinuous, especially when considering dif¬
ferences between recognized Sarracenia
species. Many individuals of one S. rubra
ssp. can look very much like to identical
to a few individuals of another ssp. in
disjunct ranges perhaps hundreds of
kilometers distant.

4) S. rubra ssp. jonesii has a range limited
to certain mountain counties in western
NC and SC (see range maps in various
references as mentioned above). Again,
small ecologic variants of the ssp. or earlv
spring pitchers have led some authors
to believe that ssp. rubra and ssp .jonesii
grow together, testimony to the confusion

Volume 11 • June 1982

41

1) Flowers of S. rubra ssp. rubra , typical of the
entire species, the only significant variation be¬
ing somewhat larger size in larger subspecies.
Note deep red petals, recurved sepals in ma¬
ture flowers. Fragrance is identically sweet or
pleasant.

3) S. rubra ssp. rubra , summer pitchers in Lex¬
ington Co., SC. Plants in this open, sunny con¬
stantly wet seep area are more robust.

2) S. rubra ssp. rubra , early spring pitchers in
eastern North Carolina. Note relatively small¬
er size of these early season pitchers in a less
than ideal habitat that will dry during the sum¬
mer.

4) S. rubra ssp. jonesu, mature pitchers. These
are taller than ssp. rubra with wider tops but
narrowing rapidly toward the pitcher base.

42

Carnivorous Plant Newsletter

5) S. rubra ssp. gulfensis in central west Florida
panhandle. While morphologically similar to
Fig. 4, the pitcher top is relatively less wide,
and of course the populauons are widely dis¬
junct.

7) S. rubra ssp. wherryi. Morphologically similar
to Fig. 6, but disjunct and in all respects uni¬
formly smaller in proportions.

6) S. rubra ssp. alabamensis. Limited (so far) to
three counties just north of Montgomery, AL,
the ssp. has a stockier, more robust pitcher
with large hoods having undulate margins.

8) S. rubra subspecies, moderately mature pitch¬
ers in comparison photo. A) ssp. rubra, B) ssp.
gulfensis , C) ssp. jonesii , D) ssp. wherryi, E) ssp.
alabamensis. (Plants in cultivation.)

Volume 11 • June 1982

43

caused by seasonal and local morpho¬
logic variations in pitchers. Generally,
the pitchers are quite tall in mature plants
(see author’s references for actual mea¬
surements) with a widely expanded pitch¬
er mouth that narrows rapidly toward the
base. The lid is well developed with a
more prominent column than in ssp.
rubra, and the upper portion of a mature,
large pitcher has somewhat of a belly¬
like bulge when viewed from the side.

5) S. rubra ssp. gulfensis, limited to a
small range in central western Florida
panhandle where it is disjunct from other
S. rubra sspp. Superficially similar in ap¬
pearance to ssp. jonesii, the top is less
wide, narrows less rapidly towards the
base, and the profile bulge in the upper
pitcher is absent to slight. Again, the
presence of small individuals, often in
less than ideal habitat has led some work¬
ers to feel that the ssp. is identical to
ssp. rubra. An additional factor to con¬
sider in this and the other Gulf coastal
sspp. is the problem of hybridization and
backcrossing with other Sarracenia spe¬
cies. Considerable experience and insight
may be required to sort these out in the
field!

6) S. rubra ssp. alabamensis so far seems
to be limited to three counties located
just north of Montgomery, AL. The pitch¬
er proportions and contour of this ssp. and
the next seem relatively more different
as a small group than the three preceding
sspp. The pitcher is stockier in appear¬
ance although the plants grow nearly as
tall as ssp. jonesii. The top is wider and
narrows very gradually to the base in
mature summer pitchers, although spring
pitchers are very similar to other sspp.
The very large hood is the most markedly
undulated (wavey) on the margins of all.
The pitcher also tends to a more pale
green background color in moderate
light as compared to the tan-green color
of the preceding sspp., although in full
light this is less apparent. A tendency to
fenestrations (alveolae, light windows,
etc.) is more obvious in this ssp. than
others, although by no means are they
clear-cut or as obvious as some other

Sarracenia species. I feel they may seem
more apparent due to the paler back¬
ground color. Similar alveolae can be
seen in the anthocyanin free variant of
S. rubra ssp .jonesii (all yellow-green pitch¬
ers, yellow flowers) which have been
found in two locations so far (5).

7) S. rubra ssp. wherryi , the last ssp.
1 recognize, is disjunct in extreme south¬
ern Alabama but north of Mobile Bay,
particularly in Baldwin and Washington
Cos. Here, the ssp. occurs with other
species of Sarracenia (no other species
have been found yet occurring with ssp.
alabamensis above) and one must be
cautious about hybrids, as in ssp. gulfen¬
sis mentioned in 5 above. The pitcher
appears similarly proportioned to ssp.
alabamensis but overall measurements
are shorter, slightly narrower, and there
is more red pigment on the average in
most pitchers.

8) Finally, we come to a comparison
photo of the pitchers of all five sub¬
species made from cultivated pitchers.
The pitchers are more mature than the
very pleomorphic, often non-specific
spring ones, being early summer, but
not as mature as full summer pitchers
illustrated in previous photos. This mid¬
dle stage is purposefully shown here to
complete the spectrum. While it illus¬
trates the differences between sspp. fairly
clearly, it also illustrates a stage when
many field observations may be made
in late spring to early summer. The letter
keys are given in the legend.

This has been the most difficult of the
primers to present in our space limita¬
tion because here in Sarracenia rubra the
differences felt to be present are based
more on degree of a character manifesta¬
tion than on a simple presence/absence
factor as has been the case with most
of the variants in the preceding two
primers. I would reemphasize that in the
case of S. rubra especially, one should
consult at least some of the technical
literature where important details, ad¬
ditional characters and measurements
are presented along with more illustra¬
tions.

44

Carnivorous Plant Newsletter

SELECTED REFERENCES

1) BELL, C. R 1949. A cyto taxonomic study
of the Sarraceniaceae of North America.
J Elisha Mitchell Sci. Soc. 65:137-166,
PI. 8-14.

2) CASE, F. W. and R. B. CASE. 1974.
Sarracenia alabamensis, a newly recognized
species from central Alabama. Rhodora
76:650-665.

3) CASE, F. W. and R. B. CASE. 1976. The
Sarracenia rubra complex. Rhodora 78:270-
325.

4) McDANIEL, S. 1971. The genus Sarracenia
(Sarraceniaceae). Bull. Tall Timbers Re¬
search Station (Tallahassee, FL), No. 9.

5) SCHNELL, D. E. 1977. Infraspecific vari¬
ation in Sarracenia rubra Walt.: Some ob¬
servations. Castanea 42: 1 49- 1 70.

6) SCHNELL, D. E., 1978. Sarracenia L. petal
extract chromatography. Castanea 43: 107-
115.

7) SCHNELL, D. E. 1978. Systematic flower
studies of Sarracenia L. Castanea 43:211-
220.

8) SCHNELL, D. E. 1978. Sarracenia rubra
Walter: Infraspecific nomenclatural cor¬
rections. Castanea 43:260-261.

9) WHERRY, E. T. 1929. Acidity relations
of the Sarracenias | Wash. Acad. Sci.
19:379-390.

10) WHERRY, E. T. 1972. Notes on Sarracenia
subspecies. Castanea 37: 1 46- 1 47.

SPECIAL NOTICE

If you have questions regarding your
subscription, please do NOT call the Ar¬
boretum office. CPN uses the Arboretum
address for mail, but the Arboretum staff
is not involved with any aspect of CPN.
Please send any inquiries by letter; your
questions will be answered promptly if
you enclose a SASE.

Occasionally, as in the case of the
March issue, printing difficulties cause
an issue to come out later than usual.
Please be patient. If it appears that your
issue was lost in the mail, we will send a
replacement as usual.

Reprints of Volumes I-IV are still un¬
available. Please do not send orders for
these volumes until notice of availability
appears in CPN.

A Miscellaneous List of Places
Where CPN is Cited

By L. Mellichamp

Shetler, S. G., “Carnivorous Plant News¬
letter, ’’TAXON 27, 478. 1978.

Slack, Adrian, Carnivorous Plants, MIT
Press, 1979.

Schnell, D. E., Carnivorous Plants of the
U.S. and Canada, John F. Blair, Pub¬
lisher; 1976.

Schnell, D. E. & D. W. Kidder, Cluster
Analysis of the Genus Sarracenia L. in
the Southeastern United States. Cas¬
tanea 41: 165-176. 1976.

The Kew Record of Taxonomic Literature
for 1974, London. Darlingtonia and
Sarracenia.

Schnell, D. E., Infraspecific variation in
Sarracenia rubra Walt.: Some observa¬
tions. Castanea 42: 149-170. 1977.

Schnell, D.E., A critical review of pub¬
lished variants of Sarracenia purpurea
L. Castanea 44:47-59. 1979.

Schnell, D. E., Notes on Utricularia sim-
ulans Pilger (Lentibulariaceae) in
Southern Florida. Castanea 45: 270-
276. 1980.

Case, F. W. & R. B. Case, The Sarracenia
rubra complex. Rhodora 78: 270-325.
1976.

N & V (Continued from page 31.)

calculate the prodigious numbers of small
insects trapped by one large bushv plant.
He placed small bits of beef on some
experimental plants and “in some cases”
the pieces disappeared. He further ob¬
served that trapped insects lived but a
short period of time although often held
by as few as one to four hairs. So, you folks
living near good patches of Proboscidea
look into this and let us know. At worst,
you could end up with some interesting
pickles.

DES

Volume 11 • June 1982

45

CULTIVATING THE ORCHID
FLOWERED BUTTERWORTS

By Steve Smith Rd #1, Box 296 Kirkwood, NY 13795

Cultivating the orchid flowered but-
terworts can be a very rewarding exper¬
ience. The rewards include delicately
shimmering leaves, beautiful flowers
in the spring and sometimes leaves on
plants ranging in size from 4 cm (P. cy-
closecta ) to 20 cm (P. moranensis ). The six
species of Pinguicula that form the Orcheo-
santhus section are: P. colimensis, P. cy-
closecta, P. gypsicola, P. macrophylla, P.
moranensis and P. oblongiloba. I have grown
all of them with the exception of P. ob¬
longiloba, which should be added by the
spring of 1982. My collection also in¬
cludes other species that respond well
to the same cultural conditions as do the
orchid flowered butterworts. These in¬
clude P hlacina (whose leaf margins roll
down instead of up), P. parvifolia and
P. hirtiflora.

These plants respond exceptionally
well to cool, damp conditions t fiat can
be provided in a building cellar. Ideal
temperatures for good growth will be a
maximum of 26°C (78°F) during the
summer and down to 10°C (50°F) during
the winter dormancy period. Do not ex¬
pose to freezing temperatures as you will
surely lose your plants. Day and night
temperatures should vary only by a few
degrees for best results. The humidity
is generally between 60 and 90 percent
during the summer months and drops
between 40 and 50 percent during the
winter months, and appears to be ideal
for these plants.

Having experimented with natural
and artificial lighting, best results were
obtained using the latter. I use a four-
tube fluorescent fixture consisting of
two Verilux Tru-bloom bulbs, one cool
w'hite bulb, and one Sylvania Gro-lux
bulb. Placing the plants from 30 to 38 cm
below the lights will give good size and will
enhance good coloration. Distance can be
increased to 61 cm or more which will

result in larger plants, however, longer
distances w ill cause loss of delicate color¬
ation of the leaves.

I have experimented with several
growing mediums, and to date the best
results have been obtained using chopped
sphagnum moss. Plants grown in peat/
vermiculite mix did reasonably well, as
did plants grown in pure peat. The
sphagnum is chopped into approximate¬
ly 3 cm long pieces and lightly packed
in the bottom of a 7.5 cm shallow pot to
within 2 cm of the top. Then the pot is
filled to the top with milled living sphag¬
num, which I chop in an ordinary food
blender. The milled top layer prevents
the larger pieces of sphagnum from over¬
growing small plants and dormant buds.
One plant is planted in each pot ex¬
cept for P. gypsicola (four per pot), P.
cyclosecta (three or four per pot), and P.
parvifolia (two or three per pot). At this
point I would like to add that despite
reading several articles written by other
growlers on adding limestone to their
growing medium, my plants were grown
without any lime whatsoever. Plants were
extremely healthy and prolific, indicating
lime is not a necessity for healthy growth.

(Continued on page 51.)

Two forms of P moranensis:

P. moranensis Oaxaca (left)

P X kewensis

Photo by S. Smith

46

Carnivorous Plant Newsletter

P. cyclosecta P. lilacina

Photos bv S. Smith

' *

I

P. moranensis

P. paruifolia

Volume 11 • June 1982

47

Flowers of D. stolonifera
“Alpine form”

Allen Lowrie looking at colony
of D. stolonifera “Alpine form”
pink flowers.

Habit of D. stolonifera “Alpine form”

48

Carnivorous Plant Newsletter

A FIELD TRIP TO TOOLBRUNUP PEAK*

Allen Lowrie, 6 Glenn Place, Duncraig, 6023, Western Australia

In November 1981, after my friend
Robert Oliver had returned from a week¬
end with his family in the Stirling Ranges
(Albany region), he reported to me that
on the cliffs of Toolbrunup Peak, at about
the 700M mark, he spotted what he
thought might be a C.P. in the distance.
He didn’t really take much notice because
the plants were off the main track up
the mountain face. After talking about
this C.P. possibility, we decided to or¬
ganise a field trip to Toolbrunup Peak
to investigate. I was rather excited at the
possibility of finding a new species, since
to date no Western Australian C.P. had
been discovered at this altitude. (The
Stirling Range area has the highest range
in the southwest of Western Australia.)

On a Friday night in November 1981,
Robert and I set off on the 400 KM
drive to the Stirling Range. We arrived
about 1:30 a.m. and crashed for the
night under the stars. At first light, after
breakfast, we set off for the walk up the
mountain. The first 300M was okay as it
consisted of slowly rising trails; after¬
wards, however, it was straight up. We
were mostly on all fours climbing up and
over large rocks, tree stumps and what-
have-you. By stopping every 50M or so
and resting, we finally arrived at the
700M mark. Walking off the main track
to the right and slowly working our way
up the cliff face we arrived at the place
where Robert had spotted the C.P. Sure
enough, they were C.P., but I couldn’t
identify them immediately. On further
investigation, I concluded that they were
a form of D. stolonifera. There are four
basic growth forms of D. stolonifera found
generally, but this one was a different
form altogether, no doubt an alpine form,
as in this region snow can be found on
Bluff Knoll ( 1073M). Toolbrunup Peak
height is 1052M.

‘Albany Region, West Australia

After crawling up the face of Tool¬
brunup Peak between the 700M mark
and the summit, I was lucky enough
to find plants in flower. What a surprise
to find these D. stolonifera “Alpine form”
sporting pink flowers! Down on the lower
sand plain, D. stolonifera is always found
with white flowers. The trip up the moun¬
tain had certainly produced a new variety
of C.P. D. stolonifera “Alpine form-Pink
flowers” was trulv the highlight of all my
1981 field trips. After Robert and I had
explored the upper face of Toolbrunup
Peak, we called it a day and set off down
the mountain. I don’t know which was
worse — the climb up or the jarred legs
on the way down.

Since it was November, the sand plain
D. stolonifera had finished and were dy¬
ing back and making up their new tubers,
but the D. stolonifera “Alpine forms” were
just coming into flower on Toolbrunup
Peak. Where the D. stolonifera “Alpine
form” grows is constantly wet, in com¬
plete shade all day long, as the plants
were found on the South side of the
mountain only. The plants were also
found mostly growing under rock ledges
and in a few cases we found groups of
plants in small caves approximately 1M
back into the cliff face. Even if the sun
did get over to the south side of the
mountain, the plants would be in com¬
plete shade all dav long anyway. This
type of C.P. discovery is fairlv common
here in Western Australia, new varieties
are always coming to light anytime one
can find a different soil type or differ¬
ent microclimate. Over the vears I have
only explored the one altitude (the sand
plains and hills). The discovery of D.
stolonifera “Alpine form-Pink flower” at
the top of Toolbrunup Peak leads me to
believe that the other peaks in the Stirling
Range may produce different forms and
species of C.P.

Volume 11 • June 1982

49

Review of Recent Literature

Barrie, N. and M. Honda (photos). 1982.
Those blooming meat eaters. National
Wildlife 20:40-45.

A popular article on Sarracenias with
very brief introductory text, but with
six color photos, some two page spread.
The photos include the obligatory
“view from (he pit,” of course. Mod¬
erately interesting.

Disney, R. H. L., Megaselia corkerae, a new
species from Nepenthes in Hong Kong
with reevaluation of the genus En-
donepenthia (Diptera: Phoridae ). Orient.
Insects 15 (2): 200-206. 1981.

The author found that these insects
breed in the pitcher fluid of N. mirabilis.

Robertson, A. and B. A. Roberts. 1982.
Checklist of the alpine flora of Western
Brook Pond and Deer Pond areas, Gros
Morne National Park. Rhodora 84: 101-
1 15.

Among the plants listed in the diverse
habitats of this Newfoundland, Canada
coastal montane area are the following
CP species: Sarracenia purpurea, S. pur¬
purea, f. heterophylla, S. purpurea v. venosa
(sic), Drosera rotundifolia (inch a f. bre-
viscapa), D. intermedia, Utricularia minor,
U. intermedia and Pinguicula vulgaris.
A profile map and brief description
of the varying habitats are given. This
is a first published location for S. pur¬
purea f. heterophylla in this area. Note
the unlikely and mistaken “v. venosa"
under S. purpurea.

Sibaoka, T. 1980. Action potentials and
rapid plant movements. In PLANT
GROWTH SUBSTANCES 1979 (F.
Skoog, editor), Springer-Verlag, New
York. Pp. 462-469.

Mimosa and Dionaea are discussed,
along with some measurements and
comments, with results similar to those
presented by Williams, Pickard and
others in the past. Aldrovanda is new
work reported in the paper in which
stimuli were applied and sequential

measurements made via insertion of
microelectrodes. New figures and ob¬
servations on speeds, direction and
extent of action potentials are pre¬
sented. The mechanism is somewhat
similar to Dionaea except for numer¬
ical differences, and Adrovanda usually
only requires one sensory hair stim¬
ulus to initiate closure.

Sutherst, R. W., et. al. 1982. Tropical
legumes of the genus Sytlosanthes im¬
mobilize and kill cattle ticks. Nature
295:320-321.

While the genus has not been demon¬
strated to be carnivorous per se, it
is of interest that tick larvae, which
crawl to the tips of vegetation to await
an animal host, are both immobilized
in the glandular secretions of some
species of this genus, and then killed
bv an aromatic toxin as demonstrated
bv partition experiments. This prop¬
erty as well as the nutritious character
of this legume may prove of value in
animal husbandry.

WhiteselhJ- K , Matthews, R. S., Minton,
M., Sc A. M. Helbling. Total Synthesis
of Sarracenin. J. Am. Chem. Soc. 103
(12): 3468-3472. 1981.

The authors described the synthesis
of the iridoid monoterpene sarracenin
from Sarracenia species.

Williams, S. E. and B. G. Pickard, 1980.
The role of action potentials in the
control of capture movements of Dros¬
era and Dionaea. Springer-Verlag, New
York. Pp. 470-480.

This and the paper by Sibaoka reviewed
elsewhere in this section were read as
part of 10th International Conference
on Plant Growth Substances held in
Madison, WI, 22-26 July, 1979. This
paper is an excellent review of the
author’s work on action potentials in
the genera and is a very useful sum¬
mary of the several papers published
by the authors over the years.

50

Carnivorous Plant Newsletter

Small P. gypsicola culling

Photo by S. Smith

Orchid Butterworts (Continued from p. 46.)

With the exception of P. hirtiflora and
P. lilacina, these Pinguicula form dormant
buds during the winter months. Leaves
become short, succulent and numerous.
These succulent leaves are the best for
propagation, and up to % can be removed
from the central bud with no ill effects
on next spring’s growth.

As with other CP, water must be pure
for best results. When and where possible
it is best to use rain water. When leaves
overlap each other and drape over the
side of the pot, care must be taken not
to wet the leaves when watering. A large
bulb syringe is excellent for watering
as the leaves can be carefully lifted and
the plant can be watered with a gentle
stream.

During the active growing season plants
may be fertilized by lightly misting with
a Vh to l/ strength balanced fertilizer once
every four to six weeks. Fish emulsion
is not recommended as it entices fungus
growth.

These plants can be propagated either
sexually or asexuallv. Thick, succulent
winter leaves are spread on milled sphag¬
num or peat or vermiculite. The medium
is lightly moistened and pot placed in a
plastic bag that is sealed and placed in a
warm, well lit environment. Approxi¬
mately four to six weeks later small plant-
lets will begin to grow from the leaf base.
When these plants are large enough to
work with they are removed and planted
in individual pots. Flowers are not self
pollinating and pollination must be done
by hand. This procedure is the same as
for P. lutea described in CPN Vol. 8,
#2, page 58. Seed pods begin to swell
about 10 days after fertilization and
mature four to six weeks later. Seeds
are spread on milled sphagnum or peat
to germinate. Seeds take approximately
three to four weeks to germinate.

Those who are interested in obtaining
these species should contact W. I P., P. O.
Box 303, Grant, Florida 32949 during the
active growing season.

Volume 11 • June 1982

51

Pinguicula macroceras ss

p. montensis, growing in Northern California.

Photo by J urg Steiger.