June 2001 North American Native Orchid Journal

NORTH AMERICAN
NATIVE ORCHID JOURNAL

____________________________________
Volume 7 June
Number 2 2001
a quarterly devoted to the orchids of North America
published by the
NORTH AMERICAN
NATIVE ORCHID ALLIANCE
* * * * * * *
* * * * * * *
IN THIS ISSUE:
REPRODUCTIVE BIOLOGY OF THE LADY'S-
SLIPPERS.
UPDATE ON CORALLORHIZA BENTLEYI
BOOKS AND PAMPHLETS ON THE ORCHIDS OF
THE UNITED STATES AND CANADA
OUT OF THE LOOP……………………………….and
more
The North American Native Orchid Journal (ISSN 1084-
7332) is a publication devoted to promoting interest and
knowledge of the native orchids of North America. A
limited number of the print version of each issue of the
Journal are available upon request and electronic
versions are available to all interested persons or
institutions free of charge. The Journal welcomes article
of any nature that deal with native or introduced
orchids that are found growing wild in North America,
primarily north of Mexico, although articles of general
interest concerning Mexican species will welcome.
Requests for either print or electronic copies should be

sent to the editor:
Paul Martin Brown, 10896 SW 90th Terrace, Ocala, FL
34481 or via email at naorchid@aol.com.
NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 7 June
Number 2 2001
CONTENTS
NOTES FROM THE EDITOR
115
REPRODUCTIVE BIOLOGY OF THE LADY'S-
SLIPPERS.
I. Introduction and the Pink Lady's-slipper,
Cypripedium acaule Ait.
Charles L. Argue
117
UPDATE ON CORALLORHIZA BENTLEYI
BENTLEY'S CORALROOT
Stan Bentley
140
BOOKS AND PAMPHLETS ON THE ORCHIDS
OF THE UNITED STATES AND CANADA
Ronald A. Coleman
143
OUT OF THE LOOP
The Slow Empiricist
154
A NEW ZEALANDER LOOKS AT ORCHIDS IN
THE WILD WEST
Ian M. St George
159
THE ARCTIC ORCHIDS OF ADELBERT VON
CHAMISSO
Mark Nir
169
RECENT TAXONOMIC AND
DISTRIBUTIONAL NOTES FROM FLORIDA
10.
Paul Martin Brown
176
6th ANNUAL NORTH AMERICAN NATIVE
ORCHID CONFERENCE
181
NEW TAXA AND COMBINATIONS
Paul Martin Brown
186
Book Reviews:
Native Orchids of Nova Scotia
Carl Munden
187
LOOKING FORWARD:
September 2001
189
Unless otherwise credited, all drawings in this issue are by Stan Folsom
The opinions expressed in the Journal are those of the authors. Scientific
articles may be subject to peer review and popular articles will be examined for
both accuracy and scientific content.
Volume 7, number 2, pages 115-189; issued June 31, 2001.
Copyright 2001 by the North American Native Orchid Alliance, Inc.
Cover: Platanthera hookeri by Stan Folsom
NOTES FROM THE EDITOR
The 2001 orchid season is full upon us

throughout North America by now and many of our
members have already made their annual forays for new
and exciting things. The September issue will be
devoted to a series of long and sort articles about those
various trips. There is still plenty of room for your
story, so please send it in to me. If you can possibly do
it, please send your story as an attached document to an
e-mail; otherwise make sure it is double spaced so it can
be scanned. If you have color photos or slides please
send 1 or 2 to go with you story. Line drawings are also
welcome. I need this material no later than September
1.
There is no color in this issue as none of the

authors submitted any to accompany their articles.
Please take the time to carefully read through Dr.
Argue's excellent article on Cypripedium pollination and
115
especially note the extensive bibliography at the end. It
is one of the most complete on the subject I have ever
seen.
We will be in Maine through early October.

Please note the telephone number change below. This
will be a quiet summer for us, as we do not have any
trips planned, although getting ready for the conference
here in September will be busy enough. Our new book,
The Wild Orchids of Florida, is scheduled for release late in
the year so final editing and proofing is well underway.
More about that in the next issue.
Paul Martin Brown

Editor
PO Box 772121
Ocala, Florida 34477-2121
352-861-2565 October - May
PO Box 759
Acton, Maine 04001-0759
Late May - September
207-636-1107
Email: naorchid@aol.com
116
Argue: REPRODUCTIVE BIOLOGY OF THE LADY'S-SLIPPERS
REPRODUCTIVE BIOLOGY OF THE

LADY'S-SLIPPERS.
II. Introduction and the Pink Lady's-slipper,
Cypripedium acaule Ait.
Charles L. Argue
So different are lady's-slippers from other

orchids that botanists sometimes place them in a family
of their own. They are distinguished by a number of
characters, but the most conspicuous difference is the
deeply saccate, pouch-like lip of the flower, from which
the plants take their name.
This lip plays an important role in pollination.

Species of Cypripedium have flowers of a type known as
trap or semi-trap blossoms (Pijl and Dodson 1966,
Dressler 1981). The flowers temporarily imprison their
insect pollinators and force them to follow a prescribed
sequence of behaviors in order to obtain their release.
An insect of the appropriate size, usually a bee, enters
the lip through the obvious large opening at its top. The
slippery inner surface and the in-folded margins of the
lip supposedly prevent it from leaving by the same route
(Summerhayes 1951, Proctor & Yeo 1973), but Knoll
(1922) and Daumann (1968) have shown that bees are
unable to escape from the lip of the European C.
calceolus even after the in-folded margins are cut away.
117
118
Figure 1. Sketches of the flower of Cypripedium acaule. A. Side view

of flower. B. Gynostemium and staminode as seen from
inside the flower. The gynostemium is comprised of the
united style and the filaments of the two fertile anthers.
The staminode is modified from a third sterile anther. C.
Flower in partial longitudinal section showing path of
bumblebee queen. a = anther, b = bract, e = escape or
exit hole, gy = gynostemium, l = labellum, lo = labellar
opening, o = ovary, s = sepals, sd = staminodium, st =
stigma.
119
Although bees have been reported to sometimes

chew through the labellum in the showy lady's-slipper,
C. reginae Walt. (Guignard 1886) and pink lady's-
slipper, C. acaule (Stoutamire 1971), most find a
different way out. A foothold is provided by tightly
packed, long hairs (trichomes) on the bottom of the lip
(Ziegenspeck 1936, Summerhayes 1951, Stoutamire
1967, Proctor & Yeo 1973). These lead along a pathway
("haarstrassen") toward escape holes at the base or heel
of the slipper (Fig. 1). The escaping bee may also be
attracted along this path by colored lines on the lip's
inner surface (Artz 1954) and by light coming from the
escape holes or, in some species, from translucent areas
("light windows") in the side of the lip near its base
(Webster 1886, Troll 1951).
Daumann (1968) found that bees are well able to

find their way out when these "windows" are covered,
but that light gradients are nevertheless a definite
orienting factor. Thus it took a bee an average of 11
minutes to find its way out of the labellum of
European yellow lady's-slipper, Cypripedium calceolus
under natural light conditions, but only two or three
minutes when an external light source was focused on
the base of the labellum. If the light was focused on the
apex the time was increased to 30 minutes.
Nilsson (1981) believes that the hairs inside the

labellum may have an additional function. Droplets of
oil which are present on their distal tips could absorb
body odors (pheromones) from visiting bees and these
odors may serve to attract additional pollinators (see
120
below). Because these hairs are strongly light refractive,

Ziegenspeck (1936) speculated that they might also
stimulate a phototactic response complementing that of
the light coming from the base of the labellum.
In its escape the bee must pass two points where

the passageway is narrowed. At the first of these it is
forced to squeeze under and rub its back against the
surface of the stigma (Fig. 1B, C) (e.g., Stoutamire
1967). The stigma provides leverage that allows the bee
to push down on and slightly depress the elastic
labellum, thus enlarging the passageway for its escape
(e.g., Wright 1975, Nilsson 1979). Ziegenspeck (1936)
considered that the basal trichomes might reduce
friction between the base of the labellum and the insect
at this point. The second narrow passage is the exit hole
itself (Fig. 1A, C). One exit hole is located on either side
of the base of the flower. An anther is so positioned
beside each exit that a bee of the proper size cannot
force its way out without contacting the anther and
carrying away a mass of sticky pollen on the dorsal
surface of its thorax (Fig. 1B, C) (Pijl and Dodson 1966,
Stoutamire 1967). Since the bee contacts the stigma
before the anther and usually does not reverse
directions, it does not ordinarily transfer pollen to the
stigma of the same flower (functional protogyny
according to Faegri and Pijl (1971)). Rather, pollination
is effected when and if the bee, upon escaping from the
first flower, is subsequently trapped again, usually in a
different flower, and the escape procedure is repeated.
121
The same bees are often trapped more than once

in flowers of the California lady's-slipper, Cypripedium
californicum (Kipping 1971) and other species of
Cypripedium (Nilsson 1979). Davis (1986), for example,
observed five bumblebees follow the prescribed course
of entry and exit from flowers of C. acaule at a site in
Massachusetts. Two were carrying pollen masses on
their thoraxes when they entered the flower and each
deposited the pollen on the stigmas of these flowers. In
a study in Nova Scotia, O'Connell and Johnston (1998)
found a greater than 90% correlation in male and female
reproductive success. In other words, over 90% of the
flowers that had a pollinium removed also received one.
Visitation rates were low, but once removed a pollinium
had a 36% to 51% chance of being transferred to the
stigma of another flower. These and similar
observations in other taxa seem to refute the contention
that once having endured the ordeals of entrapment and
the subsequent lack of reward, bees avoid repeating the
experience (Webster 1886, Baxter 1889, but see Gill
1989).
Botanists are uncertain just why bees enter the

flowers in the first place. In some cases the entry is
inadvertent. The bees, exploring the outer surface of the
lip, tumble into the trap. This has been associated with
an inflected rim near the staminode, the so-called sliding
zone (Nilsson 1981). In other cases entry appears quite
deliberate (e.g., Nilsson 1981). Daumann (1968) believes
that insects may collect oil from the hairs on the inside
of the lip. This has yet to be confirmed. It has also been
conjectured that small amounts of nectar are present or
122
that the bees feed on the hairs in the labellum (Nilsson

1979 and references therein). But it now appears likely
that the flower provides no food, and the bees are
simply deceived by false nectar guides, color, and the
odor of the blossoms, which promise nectar where none
is available (Nilsson 1979). In addition, the staminode,
which projects into the labellum, is often bright yellow
in color with contrasting spots and may appear to be a
source of pollen (Vogt 1990).
Stoutamire (1971) believes that bees must learn

by experience which flowers offer the best nectar
reward (orchid pollen can't usually be deliberately
collected by bees because of the anthers' position on the
flower) and will avoid flowers that offer no reward at all.
According to this view seed production in the
lady's-slippers may be dependent upon naïve or possibly
"forgetful" pollinators, insects that are newly hatched,
new to the area, or shifting from a depleted to a new
food source (Delpino 1874, Pijl 1966, Dressler 1981,
Ackerman 1986).
However, Nilsson (1979) contends that, in

addition to general food deception, the floral attractions
in the European C. calceolus are attuned to other
instinctive responses in bees and that very little learning
is involved. The composition of the floral fragrance in
this orchid is uncommon, which suggests that any allure
it may have differs from the usual advertisement of a
food (nectar) source (Nilsson 1979). In addition to a
monoterpene alcohol called linalool, which may elicit a
feeding response, the orchid's fragrance contains
123
acetates and farnesene. The acetates are found in

cephalic pheromone secretions of Andrena bees, and
farnesene is a component of the abdominal Dufour
gland in female Andrenas (Nilsson 1979, Bergstrom and
Tengo 1974). Pheromones are used to odor-mark
objects which then attract bees of the same species.
Farnesene is found as a lining in the nest and also
signals the location of the nest site (Bergstrom and
Tengo 1974). According to Butler (1965), females of
Andrena flavipes deposit farnesene in the soil surrounding
the nest site, and the odor provokes instinctive landing
responses in both sexes. The opening in the labellum
may mimic the opening of the nest tunnel (Catling and
Catling 1991). In addition, as already noted, several
features of the hairs within the labellum suggest that
they may be well adapted to the absorption of
pheromones directly from visiting bees as a supplement
to the artificial pheromones produced by the orchid
(Nilsson 1979).
A survey of 8 species of North American orchids

found comparable acetates present only in the ivory
lipped lady's-slipper, Cypripedium kentuckiense and one
variety of yellow lady's-slipper, C. parviflorum (probably
var. makasin based on provenance) (Barkman et al.
1997). Contrary to Bergstrom et al. (1992), Barkman et
al. (1997) found no farnesene in variety pubescence.
Nilsson (1981) notes additional differences in the
chemical composition of the floral fragrances in C.
calceolus and variety parviflorum. These differences may
correlate with differences in primary pollinator species,
as pheromone profiles can differ intragenerically among
124
pollinators (Tengo 1979). A varying blend of odor

constituents may have evolved to stimulate an
instinctive or learned pattern of response on the part of
a range of pollinators to food, sexual reproduction, or
nest location (Nilsson 1979, Gregg 1983). Much work
remains to be done in this area, especially on American
representatives.
Among species of Cypripedium the size of the

flower, the width of the entrance, and especially the
diameter of the anther-exit holes and the space between
the labellar floor and the stigma determine the size of
the insect involved in the pollination (Stoutamire 1967,
Catling and Knerer 1980). The dorsal-ventral thickness
of the thorax appears to be of critical importance
(Catling and Knerer 1980). In addition, Nilsson (1981)
noted that the depth of the labellum in the European C.
calceolus must exceed the length of the pollinating bee by
a minimum of 3 to 4 mm or the bee can simply crawl
back out through the labellar opening. All of these floral
characters are clearly under strong selection pressure in
relation to the primary pollen vectors. Research is still at
an early stage, but the pollinators of some species of
lady's-slippers can now be tentatively identified.
The largest exit holes are found in the pink

lady's-slipper (Cypripedium acaule Ait.) (Stoutamire 1967).
This orchid can occur in clusters of two to ten plants
(Wright 1975). Vegetative propagation is by rhizomes,
and adjacent plants may be from the same genet (e.g.,
Wright 1975, Cochran 1986). A single flower with a
disproportionately large, bladder-like and pendant
125
labellum is produced on a short scape from a pair of

basal leaves (Cribb 1997, Luer 1975, Gill 1989). The
flower blooms for about three weeks but senesces in
less than six days when pollinated (O'Connell and
Johnston 1998). The labellum is pink to more or less
purple with a parallel ridge on either side of the
opening. This ridge differs in color and vein pattern
from other parts of the labellum and is basically white
marked by pink to purple lines (Stoutamire 1971). The
opening itself is a longitudinal, usually closed slit 3-4 cm
long. A pollinator must deliberately push the edges of
the lip apart in order to enter, with the opening
subsequently closing behind it (Wright 1975). Any light
coming from the base of the labellum must be largely
from the exit holes, as no translucent areas ("light
windows") are present in this species (Wright 1975).
However, the base may be lighter or may contrast in
color from the rest of the lip (Wright 1975), and the
orienting pathway of tightly packed hairs on the bottom
of the lip, referred to earlier, is present (Wright 1975).
The stigma is sticky and grooved rather than simply
papillate as in other species of Cypripedium (Luer 1975,
Cochran 1986). The lateral petals and the sepals vary
from yellow-green to greenish brown, usually with a
purple tinge (Luer 1975). Along with the labellum, they
produce a sweet odor, which can be detected 2-3 inches
away from the flower (Stoutamire 1967, 1971). Long
distance attraction is visual and is related to reflectance
of ultraviolet and blue-violet by the labellum and
staminode (Wright 1975).
126
In keeping with the size of its exit holes, this

orchid appears to be pollinated by bumblebee queens in
the spring prior to the emergence of the workers
(Stoutamire 1971), a time of the year when pollinators
are usually competing for nectar (e.g., Cochran 1986).
Although Gray (1862) and Muller (1883) basically
understood the pollination mechanism in Cypripedium
calceolus, the process in C. acaule took longer to clarify.
Correll, as late as 1950, considered the chewing or
piercing activity of insects such as beetles and flies
somehow necessary to its pollination, and Bingham
(1939) thought the exit holes were too small to
accommodate the escape of bumblebees.
The establishment of bumblebee queens as

primary pollinators is based, in part, on observations
made in a northern Michigan spruce-blueberry bog by
Warren Stoutamire (1967). After much effort, a foraging
bumblebee queen, Bombus vagans, common in the bog,
was captured in the lip of one flower. It carried pollen
of the pink (stemless) lady's-slipper on its thorax
from earlier visits to other flowers in the area. (The
loose pollen of Cypripedium may adhere more effectively
to hairy insects such as Bombus than would the viscidia
of more organized pollinia (Stoutamire 1971).) Bombus
vagans has also been collected carrying pollen of C. acaule
in New Brunswick (Plowright et al. 1980, Barrett and
Helenurm 1987). Subsequent examination of
bumblebees in the collection at Michigan State
University disclosed the apparent presence of
Cypripedium pollen on several specimens of Bombus
borealis (Stoutamire 1967). Wright (1975) notes that no
127
other Cypripedium is primarily dependent on bumblebee

queens for pollen transport. Further work has
implicated queens of additional species of Bombus and
females of Xenoglossodes and Psithyrus as possible
pollinators (e.g., Stoutamire 1967, Plowright et al. 1980,
Nilsson 1981, Davis 1986, Catling and Catling 1991).
A study in Ohio supports the role of Bombus and

possibly Psithyrus but casts doubt on the effectiveness of
Xenoglossodes sp. (Wright 1975). Four species of Bombus
and a Psithyrus female, all of about the same size, were
observed to traverse the labellum and remove pollen.
The relative sizes of the exit holes and the bees allowed
escape, but with sufficient difficulty to insure contact
between the thorax and the anther (Wright 1975).
Similarly, the hinge construction of the labellum allowed
the bees to force their way beneath the stigma, while the
opposing pressure of the labellar hinge assured contact
between the stigma and the dorsal thorax of the bees
(Wright 1975). A female Xenoglossodes sp., although
slightly smaller than the exit hole, was also observed to
remove pollen. However, the mean distance between
the labellar floor and the stigma exceeded the height of
this bee by several millimeters, and Wright (1975) did
not consider it an effective pollinator of Cypripedium
acaule.
Unlike the pollen vectors of most other lady's-

slippers, the bumblebees that pollinate Cypripedium acaule
may reverse direction after reaching the anther, causing
self-pollination (Macior 1974, Wright 1975). However,
self-pollination does not appear to occur unless
128
pollinators are present, and development of seed in the

absence of fertilization is probably absent (Davis 1986).
Fruiting success in Cypripedium and other orchids

which offer no reward is consistently much lower than
in orchids that provide nectar or that mimic plants that
do (Gill 1989 and references therein). Insects visit C.
acaule infrequently (e.g., Plowright et al. 1980, Barrett
and Helenurm 1987, Gill 1989): in some studies less
than 20% of the pollen masses are removed or flowers
pollinated or fruit set under natural conditions
(Plowright et al. 1980, Cochran 1986, Davis 1986, Gill
1989, Gill and Stoutamire 1990, O'Connell and
Johnston 1998). On the other hand, artificial
pollinations have been found to produce high fruit-set
in selfed and cross-pollinated flowers (70%-100%;
Wright 1975, Cochran 1986, Davis 1986, Gill 1989,
O'Connell and Johnston 1998). Thus, Cypripedium acaule
is highly fertile, but short-term seed production appears
to be limited by the effective activity or availability of
pollinators (e.g., Cochran 1986). A similar pattern has
been reported in other orchids that rely on deception
(e.g., Plowright et al. 1980, Dafni 1984, Davis 1986,
Barrett and Helenurm 1987, Gill 1989, Primack and Hall
1990, Nilsson 1992).
Many authors consider pollinator limitation to be

evolutionarily unstable. According to this view, natural
selection should maximize reproduction within
parameters set by the availability of resources with a
trade-off between current reproduction and future
reproductive success and/or growth (e.g., Cole 1954,
129
Williams 1966, Charnov 1982, Lloyd 1980, Waser and

Jones 1991). Various strategies to increase pollination
success to the level where resources are limiting should
be favored and should spread through the population.
In Cypripedium acaule these might include a modification
in floral anatomy that would allow autogamy to occur in
this already self-compatible species (Gill 1989).
Alternatively, flowers might develop that provide a
nectar reward (Gill 1989). Such an invader could be
expected to reverse the negative reinforcement of
repeated visitor behavior in non-rewarding flowers
(Dafni 1987, Gill 1989). Although the presence or the
artificial addition of nectar does not always mitigate
pollinator limitation (e.g., Ackerman 1986, Burd 1995,
Johnson and Nilsson 1999), it had a significant effect on
both pollinia removal (male function) and fruit set
(female function) in C. acaule (Cochran 1986). Finally,
according to Gill (1989), a delay in flowering would
increase the abundance of newly emerging, naïve
bumblebees available as prospective pollinators, but
might be ineffective due to a mid-summer reduction of
sunlight in the understory and the bumblebee's
preference for well exposed foraging sites. Gill (1989)
finds it surprising that no such modifications have
arisen. Given the abundance of C. acaule and its long
history, why has the predicted increase in individual
reproductive fitness not resulted in the spread or
fixation of such forms in any current population of this
orchid?
Data based on hand pollinations suggest that an

increase in fruit production can have a negative impact
130
on subsequent growth and reproduction (e.g., Lloyd

1980, Janzen et al. 1980, Montalvo and Ackerman 1987,
Primack and Hall 1990, Ackerman 1989, Zimmerman
and Aide 1989, Snow and Whigham 1989, Ackerman
and Montalvo 1990, Primack and Stacy 1998).
Therefore, even though fruit production within seasons
may be pollinator limited, an increase in pollination rate
might not significantly improve overall reproductive
success because such an increase could have an adverse
effect on lifetime fecundity (Janzen et al. 1980,
Montalvo and Ackerman 1987, Primack and Hall 1990,
Snow and Whigham 1989, Zimmerman and Aide 1989,
Ackerman 1989).
In Cypripedium acaule resource limitation resulted

in a decrease in both plant size and the probability of
flowering following repeated episodes of sexual
reproduction and fruiting (Primack and Stacy 1998).
However, according to Cochran (1986), the effects of
resource limitation are subordinate to pollinator
limitation in this species: two years of complete
pollination are equivalent to ten to twenty years of
normal fruit set, and long life spans and low mortality
would, in time, permit resource-depleted individuals to
resume reproduction.
Calvo and Horvitz (1990) also consider the costs

of reproduction secondary to pollinator limitation.
According to their model, increased fitness resulting
from higher levels of pollination and fruit set would
overcome the relatively low cost of fruiting. Studies of
reproduction in the orchid Tolumnia variegata were
131
consistent with these predictions (Calvo 1993). A

statistically significant reduction in future growth and
flowering was observed only in plants subjected to a
high pollination intensity treatment (viz., all the flowers
in the inflorescence were pollinated resulting in a mean
fruit set about 88 times greater than in open pollinated
plants). Simulations revealed that the production of only
a few seedlings per fruit could more than compensate
the cost of fruiting and that therefore selection for
higher levels of pollination should be favored. Calvo
(1993) believes that the low level of pollination and fruit
production frequently observed in nonautogamous
orchids may be due to a low correlation between fruit or
seed production and seedling recruitment. Selection for
increased levels of pollination would be ineffectively low
if an increase in seed production is not translated into
an increase in fitness. Under such circumstances
pollinator limitation may be evolutionarily stable (Calvo
1993).
Kull (1998) has reported observations on

seedlings in Estonian populations of Cypripedium calceolus
which are consistent with this view. In this study
seedling recruitment was limited by the availability of
suitable microsite conditions. Fruit-set, which appeared
to be pollinator limited, had no observed effect on
subsequent flower and fruit production (resource
limitation) and was uncorrelated with seedling
recruitment. It therefore had no significant effect on
population fitness. Similarly, Keddy et al. (1983)
observed a scarcity of microsite conditions suitable for
seedling establishment of the sparrow’s egg lady’s-
132
slipper, C. passerinum among a mosaic of seres on the

north shore of Lake Superior. On the other hand, Gill
(1989) reported high levels of seedling recruitment for
C. acaule in Virginia.
According to Primack and Hall (1990) and

Primack et al. (1994), Cypripedium acaule may perhaps
best utilize its reproductive resources over a protracted
time period by maximizing seed production during
periods when habitat conditions are favorable. By
routinely allocating resources toward the production of
many more flowers or ovules than mature fruits and
seeds, it could benefit from chance variation in
pollinator behavior or seasons when pollinators or
resources required for fruit maturation are present in
abundance (e.g., Lloyd 1980, Ackerman 1986, Cohen
and Dukas 1990, Burd 1994). Under this view, the
expenditure of resources over the lifetime of the orchid
may involve a strategy adjusted to both pollinator and
resource limitations (Ackerman 1986, 1989, Montalvo
and Ackerman 1987, Haig and Westoby 1988, Nilsson
1992, Burd 1994, but see Morgan 1993).
References
Ackerman, J. D. 1986. Mechanisms and evolution of food-
deceptive pollination systems in orchids. Lindleyana 1:
108-113.
Ackerman, J. D. 1989. Limitations to sexual reproduction in
Encyclia brugii (Orchidaceae). Syst. Bot. 14: 101-109.
Ackerman, J. D. and A. M. Montalvo. 1990. Short- and long-term
limitations to fruit production in a tropical orchid. Ecology
71: 263-272.
133
Artz, T. 1954. Frauenschuh und Spinne. Natur. Volk 84: 421-425.

Barkman, T. J., J. H. Bearnan, and D. A. Gage. 1997. Floral
fagrance variation in Cypripedium: Implications for
evolutionary and ecological studies. Phytochemistry 44(5):
875-882.
Barrett, S. C. H. and K. Helenurm. 1987. The reproductive
biology of boreal forest herbs. I. Breeding systems and
pollination. Canad. Jour. Bot. 65: 2036-2046.
Baxter, W. 1889. Fertilization of Cypripedium calceolus. Pharm. Jour.
Trans. 20: 412.
Bergstrom, G. and J. Tengo. 1974. Farnesyl and geranyl esters as
main constituents of the secretion from Dufour gland in 6
species of Andrena (Hymenopter, Apidae). Chem. Scr. 5:
28-38.
Bergstrom, G., G. Birgersson, J. Groth, and L. A. Nilsson.1992.
Floral fragrance disparity between three taxa of lady's-
slipper Cypripedium calceolus (Orchidaceae). Phytochemistry
(Oxford) 31(7): 2315-2319.
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83-139.
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134
Catling, P. M. and V. R. Catling. 1991. A synopsis of breeding

systems and pollination in North American orchids.
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Catling, P. M. and G. Knerer. 1980. Pollination of the Small
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438.
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Cochran, M. E. 1986. Consequences of pollination by chance in
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geitonogamy and evolution of deception in orchids.
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study of Cypripedium passerinum Rich. (Sparrow’s Egg Lady-
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126(1-2): 27-38.
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serial adjustment of maternal investment during one
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137
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pink lady's slipper orchid: A four-year experimental study.
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138
Wright, L. W. 1975. The pollination ecology of Cypripedium acaule

Ait. (Orchidaceae). M.S. Thesis. University of Akron,
Akron, Ohio.
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Schroter, Lebensgeschichte der Blutenpflanzen Mitteleuropas.
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Zimmerman, J. K. and T. M. Aide. 1989. Patterns of fruit
production in a neotropical orchid: pollinator vs. resource
limitation. Amer. Jour. Bot. 76: 67-73.
Charles L. Argue, Department of Plant Biology, College of

Biological Sciences, 220 Biological Sciences Center, 1445 Gortner
Avenue, St. Paul, MN 55108-1095. email: argue001@umn.edu
Dr. Argue is a regular contributor to the Journal and has written
papers on the pollination of Listera cordata, Pogonia ophioglossoides,
the yellow fringed orchis complex, and many other native orchid
species.
139
Bentley: UPDATE ON CORALLORHIZA BENTLEYI
UPDATE ON CORALLORHIZA
BENTLEYI J. V. FREUDENSTEIN,
BENTLEY'S CORALROOT
Stan Bentley
Since press time for Native Orchids of the Southern

Appalachian Mountains (June 2000), several discoveries
have greatly expanded our knowledge of Corallorhiza
bentleyi, Bentley's coralroot.
Until August 2000, this new species was known

only from one location, which is in Monroe County,
West Virginia. I first discovered the plants in capsule in
the autumn of 1996. There were only four capsules -
two from the 1996 season and two from the previous
year. This singular population has grown in number of
plants seen each subsequent season. In 1997, there were
eight plants found. But that number jumped to twenty
plants in 1998, twenty-seven in 1999, and forty-one
plants in mid-July 2000.
On August 19, 2000, I returned to a site in Giles

County, Virginia where I had, the previous year, found
some unusually tall plants of autumn coralroot,
Corallorhiza odontorhiza. In the area, I discovered forty-
140
four plants of Bentley's coralroot. This discovery

established the species in Virginia and, at the same time,
doubled the then known population for the entire
species. This Virginia site is some eight miles removed
from the original discovery site and is approximately
eight hundred feet higher in elevation. A mile from the
above site, I discovered another new Virginia site, this
one with fourteen plants.
On August 26, 2000, friends from Pittsburg,

Clete Smith and Scott Shriver, and Doug Jolley from
West Virginia accompanied me on another search for
more populations of Bentley's coralroot. A completely
new area was found to contain nearly sixty plants. Jolley
spotted the first plants (large, healthy beauties). While
still in Giles County, Virginia, this area consisted of
several small sites scattered over roughly a half-mile of
forest roadside. The most exciting discovery was several
plants in more than one site that had flowers with red
lips rather than the typically described yellow.
Searching later in September 2000, I discovered

three more Giles County, Virginia sites. Also found was
a new site in Monroe County, West Virginia which had
twenty-nine large healthy plants. Fortuitously, this new
West Virginia site also contained plants of oval ladies'-
tresses, Spiranthes ovalis, a rare species in this part of the
country. This find is a Monroe County record.
At present then, there are six known Virginia

sites and two known West Virginia sites for Bentley's
coralroot. The total population count for the year 2000
141
exceeded two hundred individual plants. Adding to the

excitement also is several promising dried capsules
found in this past winter of 2000-2001. I am most
eagerly awaiting the arrival of the 2001 flowering season
for Bentley's coralroot.
Stan Bentley, 1201 MacGill St. Pulaski, VA 24301

Stan is a well-known native orchid enthusiast and photographer as well as the
author of Native Orchids of the Southern Appalachian Mountains published by the
University of North Carolina Press in 2001.
Corallorhiza bentleyi
Bentley's coralroot
142
Coleman: BOOKS AND PAMPHLETS
BOOKS AND PAMPHLETS ON THE

ORCHIDS OF THE UNITED STATES
AND CANADA
Ronald A. Coleman
The search for native orchids is a worthy goal,

whether for personal or scientific purposes. This
fascination, even addition to wild orchids that we share
is not a recent phenomenon. A valuable historic records
remains because many orchid students of the past took
the time to capture for posterity the fruits of their labor
in the form of articles, pamphlets and books. For those
with access to those sources, a wealth of knowledge
awaits. Collecting orchid literature can also be a passion,
and is a worthwhile adjunct to the search for plants in
the wild.
Almost from the time I started studying wild

orchids I’ve also been searching for and collecting
literature on them. The orchid literature is immense, and
it would be an expensive hobby obtaining all the orchid
books published each year, much less trying to obtain
rare and out of print publications. Just as I’ve limited
my study of wild orchids to those in United States and
Canada, so too I specialize in the acquisition of books
from that region. The list below includes all the books
143
and pamphlets on wild orchids of the United States and

Canada that I have been able to identify. All but two are
in my personal collection.
Many articles and papers on native orchids

appear each year in lay and scientific publications.
Keeping track of them is beyond the scope of this
article, although many papers are referenced in the
bibliographies of the books listed. However, some of
the pamphlets in the list first appeared in a scientific
journal and were subsequently issued as stand-alone
documents.
The publications in the list are segregated based

on their geographic coverage. Those classified as
National in scope cover all or major portions of the
United States and Canada. Those classified as Regional
cover areas equivalent to one or more states or
provinces. Those classified as Local cover a portion of a
state or province, or a specialized area such as a specific
park or district. Some of these publications are fairly
technical and are classified as Scientific. Two books did
not seem to fit any of the previous categories and are
called Special.
Some of these books, especially the regional

treatments, are still in print, and more are being added.
It is gratifying to see the number of native orchid books
covering a state or region that have been published in
the last decade, with others in press or planning stages.
Books still in print can be ordered by most bookstores.
An excellent source of in-print books is the American
144
Orchid Society. The AOS BookShop can be reached via

the internet at orchidweb.org.
Out of print books are more of a challenge, but

multiple resources are available. Two that I often use
are Raymond M. Sutton Books reachable via email at
suttonbks@getonn.net; and McQuerry Orchid Books at
www.mcquerryorchidbooks.com. The EBAY book
auction often has out of print orchid books. Go to the
EBAY web site at EBAY.com and search for book
using the wildcard orchi* as the title. Several out of
print books have been re-issued. In those cases the
dates and publishers of the original and reprint are in
the table.
Should the reader know of any books covering

the orchids of the United States and Canada that are not
in the below list, please contact the author.
Author Year Title Publisher Coverage
Ames, O. 1924 An enumeration of American National

the Orchids of the Orchid
United States and Society
Canada
Ames, O. 1910 The Genus Merry- National

(1979) Habenaria in North mount
America (reprint) Press;
(Earl M.
Coleman)
Correll, 1950 Native Orchids of Chronica National

D. S. (1972) North America Botanica
Company
145
(Stanford
University
Press)
Luer, C. A. 1975 The Native Orchids New York National

of the United States Botanic
and Canada Garden
Petrie, W. 1981 Guide to the Hancock National

Orchids of North House
America
i
i Williams J. 1983 Field Guide to Universe National
G. & Orchids of North Books
Williams, A. America
E.
Ames, B. 1947 Drawings of Florida Botanical Regional

(1979) Orchids Museum
(Earl M.
Coleman)
Baldwin, H. 1884 The Orchids of New John Wiley Regional

England & Sons
Bentley, 2000 Native Orchids of University Regional

Stanley L. the Southern of North
Appalachian Carolina
Mountains Press
Bingham, 1939 Orchids of Michigan Cranbrook Regional

M. T. Institute of
Science
Brackley, F 1985 The Orchids of New Rhodora Regional

Hampshire
(Rhodora No. 849)
146
Brown, A Field and Study Orchis Regional

P. M. 1993 Guide to the Press
Orchids of New
England and New
York
Brown, 1997 Wild Orchids of the Cornell Regional

P. M. Northeastern United University
States Press
Burian, R. 2000 Native Orchids of Oregon Regional

Oregon Orchid
Society
Cameron, 1976 The Orchids of University Regional

J. W. Maine of Maine
at Orono
Case, 1964 Orchids of the Cranbrook Regional

F. W. (1987) Western Great Lakes Institute of
Region Science
Chapman, 1997 Orchids of the Syracuse Regional

William K. Northeast University
Press
Coleman, 1995 The Wild Orchids of Cornell Regional

R. A. California University
Press
Donly, 1963 The Orchids of Regional

J. F. Nova Scotia
Ettman, 1979 An Annotated (self Regional

J, Catalogue and published
Distribution pamphlet)
147
Account of the
Kentucky
Orchidaceae
Fuller, A 1933 Orchids of North Regional

Wisconsin American
Press
Gibson, W. 1905 Our Native Orchids Doubleday Regional

H. Page and
Co.
Gupton, O. 1986 Wild Orchids of the University Regional

W. & Middle Atlantic of
Swope, F. States Tennessee
C.
Henry, L. 1975 Western Carnegie Regional

K., Werner Pennsylvania Museum
E. Buker, Orchids of Natural
and D. L. History
Pearth reprint of
Castanea
Special
Issue,
Homoya, 1993 Orchids of Indiana Indiana Regional

M. A. Academy
of Science
Keenan, P. 1983 A Complete Guide DeLorme Regional

E. to Maine’s Orchids Publishing
Co.
Liggo, Joe 1999 Wild Orchids of University Regional

and Ann Texas of Texas
Orto Liggo Press
148
Long, J. C. 1970 Native Orchids of Denver

Colorado Museum Regional
of Natural
History
Luer, C. A. 1972 The Native Orchids New York Regional

of Florida Botanic
Garden
Morris, F. 1929 Our Wild Orchids Charles Regional

& Eames, Scribner’s
E. A. Sons
Munden, C. 2001 Native Orchids of University Regional

Nova Scotia College of
Cape
Breton
Press
Niles, G. G. 904 Bog Trotting for G. P. Regional

Orchids Putnam’s
Sons
Nylander, 1935 Our Northern Regional

O. O. Orchids
149
Szczawinski 1975 The Orchids of British Regional

A.E. British Columbia Columbia
Provincial
Museum
Sheviak, C. 1974 An Introduction to Illinois Regional

J. the Ecology of the State
Illinois Orchidaceae Museum
Slaughter, 1993 Wild Orchids of Slaughter Regional

C. R. Arkansas
Smith, W. 1993 Orchids of Minnesota Regional

R. Minnesota Depart-
ment of
Natural
Resources
Smreciu, 1989 A Guide to the University Regional

E.A. & R.S. Native Orchids of of Alberta
Currah Alberta Devonian
Botanical
Garden
Summers,
B. 1981 Missouri Regional
(1987) Missouri Orchids Dept. of
Conserva-
tion
Wallace, M. 1951 The Orchids of University Regional

E. Maine of Maine
(earlier version of at Orono
above)
150
Whiting, R. 1986 Orchids of Ontario Canacoll Regional

E. & Founda-
Catling, P. tion
M.
Winterringer, 1967 Wild Orchids of Illinois Regional

G. S. Illinois State
Museum
Ames, O 1904 A Contribution to E. W.

Our Knowledge of Wheeler Local
the Orchid Flora of
Southern Florida
Bruce-Grey 1997 The Orchids of Stan Local
Plant Bruce and Grey Brown
Committee
Busswell, 1945 Native orchids of Local

W. south Florida Bulletin of
the
University
of Miami
Craighead, 1963 Orchids and Other University Local

F. C. Air Plants of of Miami
Everglades National Press
Park
Fisher, R. 1980 The Orchids of the Fisher Local
M. Cypress Hills
Fultz, F. M. 1928 Lily, Iris, and Spanish Local

Orchid of Southern Institute
California Press
151
Lamont, E. 1996 Atlas of the Orchids Torrey Local

E. of Long Island, New Botanical
York Club
reprint
Reddoch, 1997 The Orchids of the Canadian Local

Joyce M. Ottawa District Field-
and Allan (Canadian Field- Naturalist
H. Reddoch Naturalist 111(1) )
Allen, Carol 1996 North American North Scientific

ed. Native Terrestrial American
Orchids Native
Propagation and Terrestrial
Breeding Orchid
Confer-
ence
Brower, A. 1977 The Prairie White Maine Scientific

E. Fringed Orchid in State
Maine Planning
Office
Plaxton, E. 1981 North American Michigan Scientific

H. ed Terrestrial Orchids Orchid
Society
Sheviak, C. 1982 Biosystematic Study New York Scientific

J. of the Spiranthes State
cernua Complex Museum
Bulat, T. 1995 Hidden Orchids Rudi Special

Publishing
Keenan, P. 1999 Wild Orchids Across Timber Special

E. North America Press
152
Ronald Coleman, 11520 E. Calle del Valle, Tucson, AZ 85749

email: ronorchid@aol.com
Ron is the author of Wild Orchids of California from Cornell
University Press and many articles over the years in both this
journal and the American Orchid Society's publications. He is
currently working on a new book on the wild orchids of Arizona
and New Mexico.
153
Empiricist: OUT OF THE LOOP
OUT OF THE LOOP

The Slow Empiricist
Have you ever felt out of the loop when you are
trying to keep abreast of the current names and
information about native orchids? There is much new
material currently being published and researched. The
North American Native Orchid Journal has tried to keep you
abreast of this constantly changing information with
articles and news bites. It sometimes seems like a
dauntless task.
Right now there are several new publications

about to hit the bookstores that will effect your
information and knowledge about orchids. The
publishers of Flora of North America North of Mexico are
about to issue volume 26, which contains the orchids.
There are many changes in nomenclature and alliances.
If you are a relatively new orchid enthusiast you

might feel lost amidst the myriad names and labels
attached to each orchid. Especially if you are a novice,
you will need to be apprised of the procedures used to
affect these changes as well as why orchids and in fact,
all plants, have a rigorous and scientific method of
154
being identified and an appropriate name appended to

each specimen.
Let us begin with a short foray into the naming

of an orchid. When a new species is discovered it needs
a name. Science has found that common names do not
do the job as there can be too many common
appellations for a particular plant. This can result in
misapplications of names to the wrong plants or over
the years of a particular plant having several names
attached to it that are not valid. These last are called
synonyms. Since this is confusing and can result in
misidentification, science came up with a strict code
(International Code of Botanical Nomenclature) for the proper
naming of a new species. It goes something like this: the
plant has to have a Latin name. This will ensure that
people all over the world will have a common language.
There can also be affixed a common name to the orchid
in whatever language the information is being
published. There can be further information in the
name such as forms added to the description. Then the
botanist who publishes this information will have
his/her name attached to the end of the orchid's name.
This is about as simple as I can make it. There are a lot
of nuances that I have not included like a new species
being separated from an existing species cannot have a
name too close to its original name as it would be too
confusing. Believe me there are more steps and
procedures before a new species can be properly
named.
155
Maybe it would be a good idea if I took you

through the naming of a new orchid that I was privy to
last year. On an excursion along Florida's Turnpike to
count the number of Sacoila lanceolata plants that were
growing along the grassy edges of the road, Paul Martin
Brown and his cohort, Stan Folsom, discovered a new
color form (gold) for the red spiranthes. The new
name of the new color form was named for Folsom.
Therefore S. lanceolata, the red spiranthes had a new
relative, S. lanceolata forma folsomii P.M. Brown. It was
published last year in this journal and has joined the
other orchid names in the world.
The other things that have been happening all

center around advances in the scientific tools for
ascertaining what each orchid really is and in which
family it belongs. As simply put as I can describe it they
have been testing orchids for DNA and finding that
certain orchids don't belong with certain other orchids
as was originally thought.
They have also been most diligent in tracing the

origins of the names applied to the orchids to insure
that the proper designation has been supplied. Couple
these changes with the constant ongoing searches for
new species and you have a large and sometimes
bewildering array of new descriptions, or families, or
new orchids to digest and learn about.
What brought me to write this column was a

lecture I attended recently in which the speaker referred
to several of the orchid slides he was presenting by
156
names that were no longer considered correct for the

plants. Since he was a retired botanist of some years of
age it could be forgiven him that he was not cognizant
of the current appellations for those orchids.
As I stated in an earlier column, nomenclature is

governed by a strict set of rules. This publication tries to
follow the proper procedures when plant names are
used. The current researchers in the field of native
orchids are looking at all the previously cited names for
each particular orchid described to determine which
name is the earliest and correctly published name for
the orchid. That means that some commonly held
names for orchids are in error as there were previously
validly published names for these plants. Hence the
confusion when someone refers to a plant by its old,
commonly accepted name that is no longer valid.
I thought I would try to make up a list of some

of the changes that I have become aware of as I have
read about their occurrence. I'm sure that I will not have
all the current changes as they are happening nearly
every day somewhere in the world. I hope that Paul
Martin Brown will enrich the selections with more if he
has knowledge of the changes coming into effect.
The red Spiranthes, is now known as Sacoila

lanceolata. It used to be Spiranthes lanceolata. There are also
several related color forms like the green form and the
gold form.
157
The entire genus of Spiranthes has seen its ranks

swell with the inclusion of S. eatonii, which had been
hiding under the names of S. lacera var. gracilis and S.
torta. P. M. Brown discovered the plant specimens
collected by Ames in an herbarium with all the
nomenclature present but it was never validly published
until Brown published it the March 1999 Journal.
Platantheras have been very carefully analyzed by

people like Chuck Sheviak in New York State and have
had extensive changes wrought by his findings.
Epidendrum conopseum is now known as E.

magnoliae since that name was published a scant two
weeks before the other more familiar name was
published.
These are just a few of the many changes that

you all will have to become cognizant of as the new
publications come on line. I hope you will look at the
changes as a chance to enrich your knowledge and not
as an onerous chore. You wouldn't want to be called by
a misapplied name or synonym if it really didn't fit you.
Speaking for the orchids, neither do they.
Your Slow Empiricist
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St. George: A NEW ZEALANDER LOOKS AT ORCHIDS IN THE
WILD WEST
A NEW ZEALANDER LOOKS AT

ORCHIDS IN THE WILD WEST
Ian M St George
One of the books from his own childhood that my father

passed on to us was called something like Buffalo Bill’s Wild
West. It had a lot of Buffalo Bill Cody stories, and I loved them.
“With a cry, Cody sprang from the trees” and “Cody’s rifle
cracked and another redskin bit the dust” were strong fuel for a
boy’s imagination. The first western movie I ever saw was
Colorado Territory.
So the Rocky Mountains of Colorado have a nostalgic

pull for me. This is Arapaho, Pawnee and Comanche country.
Cody is buried near Denver. Wild Bill Hickock, on the other
hand, is buried at Deadwood in South Dakota. I’ve been there
too.
The “unsinkable” Molly Brown, Titanic survivor, lived in
Denver, about as high above and as far from the sea as she could
run.
Bill Jennings is a mining engineer who lives at the

foot of the Rockies in Louisville, Colorado. He has a
detailed knowledge of Colorado orchids. I phone him
from Wellington, and when I reach Colorado in early
July, he is at my hotel an hour later in his pickup (“I can
tell you’re from Colorado” he had been told on a recent
orchid jaunt to Arizona: the cracked windscreen is a
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dead giveaway, a sign of the gravel laid on Colorado

roads in winter to improve traction). We head for the
hills, through the city of Boulder and up Flagstaff
Mountain Road to an area among the red rocks and the
ponderosa pines, where we walk down a trail.
There’s a plaque on the 15th step of the State Capitol
building in Denver: it marks altitude 5280 feet: one mile.
Denver is Mile High City. Up here we are at 7500 feet,
and it makes you a little breathless. First stop is a damp
creekside where the green bog orchid, Platanthera
huronensis, is in full spike. We ascend further, pause to let
a grey/green broad-tailed hummingbird, little bigger
than a bumble-bee, take nectar from trackside flowers.
The flowers are like our New Zealand weeds, a legacy of
the miners who came and went last century. A few are
unfamiliar: the scarlet tufts of Indian Paintbrush for
instance. Further up we find giant rattlesnake orchis,
Goodyera oblongifolia, in bud, its green leaves marbled with
white. The coral-roots: spotted, Corallorhiza maculata and
Wister's, C. wisteriana, are in fruit, but by a creek is the
broad-lipped twayblade, Listera convallarioides, its
flowers translucent brownish green mothwings. Bill tells
me he has never seen a rattlesnake around here and only
once in 30 years has he seen a black bear and I am
reassured. If you come from a country with no
dangerous animals (except the domestic ones) you do
like to ask.
On the way home we stop in the Boulder

suburbs, for there between two new office carparks is
another orchid site; the developers are aware of the
treasure they guard, and have limited the size of the
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carparks to preserve it. This is Ute ladies'-tresses,

Spiranthes diluvialis, a nationally threatened orchid, and
one of them has a few white flowers fully open. I return
a few days later to find several plants in full flower.
Ten thousand acres of this forest were razed by

fire last July 4, Independence Day: why do we mark
summer celebrations with fireworks? Bill has supplied
maps and printed an itinerary for me so the next day I
am awake at 5.30, and soon on my way. Haybales in the
paddocks remind me this is summer, despite the winter
I have left behind in New Zealand. The air is so dry I
get a static electric shock every time I step out of the car
– at this time of year the relative humidity is in single
figures here. Names like Old Stage Road, Big Elk
Meadow and Beaver Meadow remind me I am in the
Wild West. Fifty miles later I am among the high peaks
in Rocky Mountain National Park. The north faces and
flat areas under steep banks still carry snowdrifts. A
Steller’s jay saunters off as my car approaches the
carrion he is eating; a chipmunk races across in front of
me. I am at the carpark near Bear Lake at 7.30, here to
look at the clustered lady's-slipper, Cypripedium
fasciculatum, a lady’s slipper with two round glossy leaves,
and a couple of brownish flowers. Alas, there has been
a mild winter, an early spring and a hot dry summer, so
they are over: I sadly photograph the spent heads. But
whoa! Beside them are what the British call the lesser
twayblade (heart leaved twayblade), Listera cordata, and
nearby, in the damp creeksides, the porcelain-white
white bog orchid, Platanthera dilatata, the yellow-
flowered P. huronensis, some other Platanthera, some
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coral-roots past flowering, and then suddenly, in full

flower, spotted coralroot, Corallorhiza maculata, its red
stems and flower parts contrasting strongly with the
white, red-speckled labellum.
Chipmunks chatter their warnings as I pass, and

squirrels scuttle up the far side of trees, away from my
camera, chirruping their distaste for my presence. In a
carpark traffic island a chipmunk delicately eats wild
strawberries, pauses, darts, pauses, darts: she sits still
just long enough for me to photograph the rock she’d
been sitting on a moment earlier. Silver birches among
the pines; spruce, aspen, juniper.
A doe feeds in the lush green of a swamp near

Cub Lake. A coyote lopes across the road, untidy as he
loses his winter coat. A marmot runs for cover under
his rocks, then bold as brass, sneaks flattened to the
ground, clear in his own mind that he is invisible, back
towards me. I take time to approach a chipmunk,
camera at eye, clicking as I come closer; I finish and rise,
smiling smugly to myself in anticipation of the
photographs, to find I am being watched by a group of
American trampers, pitying, but entertained by my
interest in such an everyday creature. Out on the flat of
the glacial moraine the members of a group of butterfly
collectors huddle, nets like triangular pennants in the
breeze, looking at their treasures. In the warmth of an
alpine summer morning the air is fragrant with the
perfume of the pines.
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I drive further up. Two elk bulls in velvet graze a

high grassland, all power and majesty. Above them the
conifers become stunted, twisted, and suddenly they are
finished and I am in open alpine meadowland, the
wildflowers colouring patches of white, yellow, orange,
pink to scarlet through crimson to mauve, purple in all
its shades, sky to dark blue to black. At the top it is
11,796 feet, a tad shorter than Mount Cook.
There is something exhilarating about treasure

hunts, and orchid forays in foreign countries often
provide that sense: “Continue east on US36 to Fish
Creek Road, go south for three miles till road curves
west, then turn south along gravel road past Fish Creek
Ranch” (It’s a “dude” ranch, my papers say
disparagingly), “enter Camp Cheley under a wooden
arch, drive a further 250 yards until you reach a point
where the power lines on your left send a branch south
across the road: there is a sign on the power pole that
reads ‘Watch out for future world leaders at work and
play’: The large yellow lady's-slipper, Cypripedium
parviflorum var. pubescens, is about 50-100 yards up the hill
on the south side of the road: there are two extra large
ponderosas right there….” (I should tell you Camp
Cheley takes young people and trains them in leadership
skills….)
Dang. The Cypripedium is also over, the remnants

of its flowers browning in the afternoon heat. Nearby
though, I am compensated with what the British call the
frog orchid, Coeloglossum viride, in full flower. At another
site I seek the flamboyant eastern fairy slipper, Calypso
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bulbosa. In 1979 I took a sabbattical leave in the

American West and it was not all work. Before I left
home I had been trying some rather pretentious
architectural photography in black and white, and when
to my surprise I found a lovely pink orchid under the
giant Californian redwoods I forgot I didn’t have colour
in the camera and took a dismal series of pictures of this
fabulous flower in shades of grey. This time I am ready
with colour film, but sadly Calypso is well in fruit by early
July.
Near Red Rock Lake I find more Listera cordata,

some plants with brown flowers, some with green. The
lake is one third covered with a beautiful native yellow
water lily, but the Indian Peaks of the Great Divide are
reflected in the stillness of the other two thirds. Bill has
mapped out a number of further stops in case I have
missed orchids, but I am replete and head back to my
hotel. Hard physical exercise is good for jetlag, but at
5pm I am flagging (it’s 11 am tomorrow in New
Zealand and it’s 12 hours since I slept).
I descend through Boulder Canyon: tanned

bareheaded denim-clad couples on Harley-Davidsons
zoom by, blonde hair flying, dangerously yet somehow
enviably unencumbered with crash helmets. A Denver
radio station plays continuous country music. Later,
down on the flatland below the foothills, prairie dogs
have turned the paddocks into a moonscape, the
entrances to their underground cities looking like
craters; one stands upright keeping watch as the others
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feed nearby. Their habitat is shrinking as the cities of

men expand.
I spend the working week sweltering in the

summer heat of the plains, and marveling at the early
“monsoon”, the evening thunderstorms that bring rain,
spectacular displays of lightning, and death. Colorado
has the second highest death rate from lightning strike
in the US: three children taking an outdoor spa bath
died recently in Boulder; mountaineers will abort their
climb if they can’t reach the summit by midday.
A week later I am on Interstate 70, heading up

into the mountains again, this time toward the great ski
resorts of Aspen, Vail and Breckenridge. There’s a dead
porcupine on the road, a bison herd grazing to the right
of the highway. The yellow tailings of gold and silver
mines streak the hillsides. Higher up a couple of stones
falling down a bank above the road catch my eye: sure
enough, two bighorn sheep are grazing near the top,
their grey fleeces perfect camouflage against the rocks.
My orchid stop is above the historic gold and

silver-mining Georgetown, its Victorian weatherboard
houses like Wellington’s. Above the town is Guanella
Pass, and the road zigzags up the hill. My guide has
written, “The road is narrow and potholed… continue
to Clear Lake and at the upper end turn left into the
parking lots. Go to the lower parking lot. The wet seepy
hillside between the parking lot and the road is full of
hooded ladies'-tresses, Spiranthes romanzoffiana, and
Platanthera huronensis.” Indeed it is, and though I spot
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the tall Platanthera quite quickly, and though I recognize

the suitability of this habitat for Spiranthes, it takes me
some time to spot the first. Spiranthes romanzoffiana is
tiny, but I spot many as I begin to get my eye in and
appreciate the tallest of them is 10cm, and most less:
5cm or so. It is vanishingly rare in Britain, reported
from only one site.
In Georgetown for the night I discover a

restaurant called “Prague”. On his menu the Czech chef
has (pause, then hushed, reverent voice), “…rack of
New Zealand lamb, 21 dollars 95”. That’s about
NZD50. I settle for pork and sauerkraut, congratulate
the chef on the food, congratulate him doubly on his
choice of a smooth Californian merlot, and tip him
generously in my postprandial largesse.
Next day I drive the highest (12,095ft)

continuous paved road in the world, called (of course)
Independence Pass. The roadside ditch west of the
continental divide is full of Platanthera dilatata at 11,000
feet. A doe, ears out and looking, makes sure it is a
camera and not a rifle I am taking from my pack, then
continues grazing. I sneak a quick look at the
pretentious tourist town of Aspen and pass on (Vail I
go past: I am told it is even worse). At Glenwood
Springs I visit the grave of “Doc” Holliday. Doc was
trained as a dentist, but when it dawned on him that
bettin’ and shootin’ were more fun than tooth pullin’ he
became a gunman and gambler. He must have been
quite good at the shootin’ for he died of pulmonary
tuberculosis.
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I have come home intending to revise my way of

helping orchid-minded visitors to New Zealand. I think
I will do what Bill did for me: guide them to orchids
that grow in wonderfully scenic sites with a set of
treasure hunt instructions, setting them free,
unencumbered by the obligation of my company, to feel
like adventurous pioneering children seeking gold in a
wild new country.
Ian St George is editor of the New Zealand Native Orchid Group

Journal.
22 Orchard St, Wadestown, Wellington, New Zealand, phone
0064 4 499 4227, fax 0064 4 3894178, email istge@rnzcgp.org.nz.
eastern fairy slipper

Calypso bulbosa var. americana
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THE ARCTIC
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Nir: ZEALANDER LOOKS ATVON
OF ADELBERT ORCHIDS IN THE
CHAMISSO
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THE ARCTIC ORCHIDS OF ADELBERT

VON CHAMISSO
Mark Nir
The purpose of this paper is to tell the story of

the origin of some orchid names encountered in the
flora of North America and of their progenitor.
The brig "Rurik," a smallish two-master of 180

tons, under the command of the Lieutenant of the
Russian Imperial Navy Otto von Kotzebue, left the port
of Copenhagen on August 17, 1815. Well financed and
for that time relatively lavishly provisioned by Admiral
Adam von Krusenstiern, one of the early
circumnavigators, it was commissioned by Count
Nikolaj Romanzoff, Chancellor of the Russian Empire,
to undertake "A Voyage of Discovery, into the South Sea and
Beering Straits, for the Purpose of Exploring a North-East
Passage." It was granted the privilege of displaying the
Russian imperial naval flag and bore a crew of thirty
sailors and eight small cannon. The first officer was
Gleb Simonovich Shishmareff; the ship surgeon Johann
Friedrich (a.ka. Ivan Ivanovich) Eschscholtz, physician,
entomologist and botanist; the painter Louis (Login
Andrejevich) Choris; and the naturalist Adelbert
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WILDOF ADELBERT VON CHAMISSO
WEST
(Loginovich) von Chamisso. The patronymics were

indispensable on a Russian ship, although of those
mentioned only Cleb Simonovich was Russian.
Chamisso was 34 years old when he joined the

expedition. He was already prominent in German
literary circles as a poet who had achieved international
fame with the novel "Peter Schlemihl," the story of a
man who sold his shadow. Born Charles Louis
Adelaide Count Chamisso de Boncourt, he was eleven
years old when during the French revolution his father,
confronted with the choice between decapitation and a
hasty exile, unsurprisingly chose the latter. Two
centuries ago life as a refugee appears to be no different
from what it is today. The family wandered for four
years in the Low Countries and in several German states
until permitted to settle in Berlin in 1796. At the age of
fifteen he helped support his family by painting
miniatures at the royal porcelain factory. His talent as a
painter is probably best attested to by the beautiful
color illustration of Eschscholzia1 californensis Cham.,
which accompanies his description of the Yellow
California Poppy, the California stare flower. Following
several years of military service and travel, Adelbert
became a student of natural history in Berlin. It was at
this point in his career that due to ill health the
naturalist originally intended for the voyage of the Rurik
had to be replaced. Through well-placed connections
and good luck Chamisso became the replacement.
1
Dr. Eschscholtz spelled his name with a "t", but Chamisso omitted it from
his new genus Eschscholzia.
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Nir: THE ARCTIC ORCHIDS WEST
ADELBERT VON CHAMISSO
The first leg of the voyage from Plymouth to

Tenerife resulted in a major scientific find: studying the
diverse marine creatures collected floating close to the
surface, Chamisso and Eschscholtz discovered the
metamorphosis of the tunicates.2
The crossing of the equator for the first time on

the 23rd of November was' accompanied by the then
customary celebrations, which nowadays arc reduced to
a terse announcement (if at all) by the pilot. They
reached Santa Cruz (off Brazil), continued southwards,
rounded Cape Horn, and after a short stay in Chile
sailed due west skirting Sala-y-Gomes, visited Easter
Island, passed between the Marquesas and French
Polynesia, then gradually turned north, revisiting
Romanzoff Island (now Wotje) in the Marshall
Archipelago, discovered by yon Krusenstiern in an
earlier expedition, and finally reached Petropavlovsk on
Kamchatka on June 19th.
The first period of exploration of the Bering Sea

and the Aleutians between July 27th and September
14th 1816, resulted in many geographic discoveries,
among them the Kotzebue Sound and in it the small
Chamisso Island, Choris Peninsula and Eschscholrz
Bay. Exploring the area provided part of the material
for Chamisso's publication in Linnaea (1828).
2
http://www.uinla6.kI2.wy.us/WWW/MS/8grnddlnlo%20Acccss/SP
ANTLGY/urochord.htm
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St. George: NEW ZEALANDER
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The Rurik then turned southwards and after a

stay in San Francisco continued towards the Sandwich
Islands. Arriving at the harbor of Hana-ruru on the
island of O'Wahu they met king Tameiameia. It is not
the purpose of this paper to summarize the rest of the
"Voyage" but the preceding sentence should serve to
whet the appetite of those so inclined, to read the whole
thing (although calling the English version a
"translation" would be too kind). A lengthy exploration
of Hawaii and the Marshalls gave Chamisso the
opportunity to make observations and measurements
published in his "Notices sur les iles de corail de Grand
Ocean" (1821), which provided much of the material
and crucial information for Darwin's "The Structure and
Distribution of Coral Reefs" (1842). The Rurik then
headed northward again, back to the Aleutians.
The second stay of the Rurik in Unalaska lasted

from April 24th to June 29th. It served mostly to
provision and equip the vessel for a second attempt to
sail farther north in search of the northeast passage, but
for Chamisso it provided the opportunity to botanize
during the period of maximum flowering. Anemones
and orchids were starting to appear towards the end of
May and flowering into June, providing material for the
rest of his Orchidaceae Arcticae.
The trip northward failed again and the Rurik

returned to Unalaska for the last time on July 22nd. The
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ship left the Aleutians on August 181h and returned

through the by then well-established route through the
Marshalls, Hawaii and Guam to Manilla, thence

to Capetown and back to Europe.
Orchidaceae Arcticae contains the following taxa (the

most informative reference follows in smaller font):
1. OrchisJ latifolia L.(?) var. beeringiana Cham. =

Dactylorhiza aristata (Fisch. ex Lindl.) Soo. Hult. Fl.
Kamtch. 259
2. Orchis camtschatica Cham. = Platanthera camtschatica
(Cham.) Makino. Hult. Fl. Kamtch. 262
3. Habenaria borealis var. albiflora Cham. = Platanthera
dilatata (Pursh.) Lindl. ex Beck. Correll, Nat. Orch. N. A. 71
4. Habenaria borealis var. viridiflora Cham. = Platanthera
convalliaefolia (Fisch.) Lindl. Hult. FI. Aleut. 151
5. Habenaria schischmareffiana Cham. = Piperia unalascensis
(Spreng.) Rydb. Hult. Fl. Aleul. 148
6. Habenaria chorisiana Cham. = Platantehra chorisiana
(Cham.) Rchb. f. Hult.. Fl. Aleut. 148
7. Habenaria viridis R. Br. = Coelogloglossum bracteatum
(Muhlenb. ex Willd.) Parl. Correll, Nat. Orch. N. A.: 114
8. Spiranthes romanzoffiana Cham. Correll, Nat. Orch. NA 220
9. Listera cordata R. Br. Correll, Nat. Orch. N.A.: 126
10. Listera eschscholziana Cham. = Listera convalllarioides
(Sw.) Torrey. Hult. Fl. Aleut. 154
11. Ca!ypso borealis Rich. = Calypso bulbosa Correll, Nat.
Orch. N.A.: 280
12. Cypripedium macranthum Sw. Hult. Fl. Kamtch. P. 257
13. Cypripedium guttaum Sw. Hult. Fl. Kamtch. P. 256
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14. Malaxis diphyllos Cham.

15. Corallorhiza innata (sphalmate intacta) R. Br. =
Corallorbiza trfiida Chat. Hult. Fl. Aleut. 156
After his return Chamisso worked as associate

curator at the botanical garden and herbarium at Berlin-
Dahlem under von Schlechtendal and later as its
director. He settled down to a rather unsettled family
life, producing offspring both marital and extramarital,
as well as a large body of poetry and a wealth of
taxonomic publications. All these before he died at age
57.
Adelbert von Chamisso drawn by E.T.A. (The

Nutcracker) Hoffmann, 1805.
References:
Darwin, C. 1842. The Structure and Distribution of Coral Reefs.
Chamisso,. A. 1819. De animalibus quibusdam e classe vermium
Lineana: De Salpa. F. Diimmler Verag, Berlin.
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Chamisso, A. 1828. Orchideae Arcticae. Linnaea 25-35. Chamisso,

A. 1975. Werke in Einem Band. Winkler Verlag, Munich.
Correll, D. S. 1978 Native Orchids of North America. Stanford
University Press.
Hulten, E. 1960. Flora of the Aleutian Islands. J. Cramer, New York
Hulten, E. 1927. Flora of the Kamchatka and the Adjacent Islands.
Almqluist & Wiksells, Stockholm
Lahnstein, P. 1987. Adelbert von Chamisso. Ullstein Verlag, Berlin.
Acknowledgment:
I thank Alfred von Krusenstiern and Dr. Frank Axelrod
for valuable help.
Mark Nir, M.D.. 18 Ashford Ave., Dobbs Ferry, NY 10522;

email: klapi@aoI.com
Mark is the author of the exhaustive Orchidaceae Antillanae
published in 2000 by DAG Media. It includes all of the orchids in
the Caribbean and many in southern Florida.
Platanthera chorisiana Malaxis diphyllos
175
Brown: RECENT TAXONOMIC AND DISTRIBUTIONAL NOTES
FROM FLORIDA 10.
RECENT TAXONOMIC AND

DISTRIBUTIONAL NOTES FROM
FLORIDA 10.
Paul Martin Brown
Just Visiting or Permanent Resident?
Florida has several species of orchids that have

been found in a wide variety of counties in the state that
are generally considered not to be part of the native
flora. In many instances that determination is clear-cut
because the first occurrence of the species can be
documented and then the spread and establishment of
the species has been noted. These species should be
classified as Naturalized: well established and
reproducing in the wild. If the original plants were
purposefully planted out in the wild they would be
considered Introduced.
In other cases the species may be a popular

garden subject and subsequently found in one or two
nearby localities, but rarely persisting. If sporadic
sightings are made then these plants could be classified
as Adventive: occurring in the wild (or at least without
cultivation) but not necessarily reproducing or
persisting.
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FROM FLORIDA 10.
The third category is a catchall for those species

that have been found but a single time, are not
reproducing and are usually short-lived. These would be
considered as Waifs.
Because of Florida's geographic position so near

the West Indies, there are several records of species
more frequently seen southward that have had one or
two Florida records. Whether to consider them as truly
native or as a casual introduction (a waif?) requires
carefully assessing the habitat in which they have been
found and were they found in areas that are infrequently
botanized.
The above categories certainly are not cast in

stone, and it is conceivable that a species that is
considered a waif or garden escape could eventually
become thoroughly naturalized. As far as some of the
Caribbean species are concerned there may never be a
satisfactory answer.
Those species that are thoroughly naturalized:
Cyrtopodium polyphyllum (Vell) Pabst ex F. Barrios

yellow cowhorn orchid
Miami-Dade & Highlands Counties; well established at
several sites
Oeceoclades maculata (Lindley) Lindley

spotted African orchid
177
FROM FLORIDA 10.
wide spread throughout central and southern Florida

and spreading northward; considered by some to be a
weed!
Phaius tankervilleae (Aiton) Blume

nun orchid
a single, well established site in Hardee County, with
large, old plants; reports from Miami-Dade County
form the 1960s
Spathoglottis plicata Blume

Javanese violet orchid
Well established in old shellpits in Palm Beach County
since the early 1980s; in many sites
Zeuxine strateumatica (Linnaeus) Schlechter

lawn orchid
the most widespread of all of these naturalized species;
throughout all of Florida and now found in the adjacent
Gulf states
Those species which have had but one or two

records and should be considered garden escapes or
waifs:
Bletia florida (Salisbury) R. Brown
slender pine-pink
as a garden escape in Homestead
Bletia patula Hooker
broad-lipped pine-pink
single collection from 1947
Bletilla striata (Thunberg) Reichenbach f.
Chinese orchid
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FROM FLORIDA 10.
As a garden escape or purposeful introduction in

Escambia County; not reproducing
Epidendrum radicans Paven ex Lindley
flame star-orchid
found at old homesites in Miami-Dade and Lee
Counties; persisting but not reproducing
Laelia rubescens Lindley
pink Laelia
a single plant of about 9 or 10 years old found in 1999
in Miami-Dade County; does not to appear to be
introduced
Vanilla planifolia Jackson ex Andrews
commercial vanilla
popular garden subject that is occasionally an escape
Vanilla pompona Scheide
showy vanilla
popular garden subject that is rarely an escape (1945)
Those species that have had few collections, by

credible collectors, and have not been subsequently
seen in the wild. All are usually considered to be part of
the native flora, but present an interesting example.
Cranichis muscosa Swartz
moss-loving Cranichis
no positive documentation since the early part of the
20th Century
Encyclia rufa (Lindley) Britton & Millspaugh
rufous encyclia
known from two collections as the same site in 1926
Lophiaris carthagenensis (Jacquin) G.J. Braem
spread-eagle orchid
179
FROM FLORIDA 10.
known form a single collection in 1916

Maxillaria parviflora (Poeppig & Endlicher) Garay
small-flowered Maxillaria
found deep in the Fakahatchee in 1975 and not
relocated in recent years despite several searches
Pelexia adnata (Swartz) K. P. J. Sprengel
glandular ladies'-tresses
found in Everglades National Park in 1977; populations
dwindled and not seen again
There are certainly several other species that could be
considered as above. It will always be a matter of
speculation by some and a drawn conclusion by others
as to the origin of those species.
Triphora gentianoides (Swartz) Ames & Schlechter, least

flowered triphora; gentian noddingcaps, presents a
peculiar problem as it has been well established since
1919 in several wild areas of central and southern
Florida, but most recently it has been found in mulch
and garden soils. Some authors consider it as native
while others see it as adventive.
Many of the above species, which are found in

southernmost Florida, have been treated in greater
detail by Roger Hammer in the March issue of the
Journal.
180
Brown: NEW TAXA AND COMBINATIONS
6th ANNUAL NORTH AMERICAN

NATIVE ORCHID CONFERENCE
September 7-9, 2001

Acton, Maine
Pre-conference workshop on Thursday evening

Sept. 6, 6:30-9 pm
Spiranthes Ecology, Distribution & Identification
Charles J. Sheviak & Paul Martin Brown
A workshop using both live and herbarium
material; microscopes will be provided and an
extensive handout will be supplied: cost $12.00
schedule
Friday, Sept. 7
9am Registration - Acton Congregational Church
Conference commences at 10 AM
Speakers will include:
Charles J. Sheviak
Mark Nir
Scott Stewart
Gerardo Salazar
Brian Keel
a special presentation from Philip Keenan on the Life
History of Triphora trianthophora
and others to be announced
181
Registration cost: $55, which includes lunch and

dinner and all field trips
Meals package for those not attending the conference:
Lunch $7.50, Dinner $10.00
Field Trips on Saturday and Sunday, September 8&9

Saturday to western Maine and nearby New
Hampshire proceed northward to Colebrook, NH
Sunday to northern New Hampshire and northern
Vermont
Species to be seen include:

Spiranthes cernua (3 different ecotypes), Spiranthes
casei, Spiranthes ochroleuca, Spiranthes
romanzoffiana, Spiranthes xborealis and several other
Spiranthes hybrids; Goodyera repens, tesselata and
pubescens (some may still be in flower); Epipactis
helleborine in a variety of colors; and the possibility
of late-flowering Platanthera lacera, psycodes,
xandrewsii, aquilonis & huronensis.
Please send your registration and indicate the various

activities and meals carefully. Unlike other
conferences, meals are included for registrants and
available for spouses or companions.
SEND REGESTRATIONS TO NANOA, PO BOX

759, ACTON, MAINE 04001
Conference Only: $55 for Friday, Saturday and

Sunday
182
Pre-conference Spiranthes Workshop: $12
Additional meals lunch on Friday $7.50

Dinner on Friday $10.00
The Acton Congregational Church is located just off

Route 109 in Acton Center and visible from the road.
It is adjacent to the Town Hall and Library just past
the new Fire Station.
From Boston take I-93 north to I-95 north to the

Route 109 exit in Maine. It is labeled for Sanford and
Wells. Take 109 north through Sanford and
Springvale, past Mousam Lake and into Acton. Past
the Acton fairgrounds; it is a small town with no
commercial center - just a crossroad
Driving time from Boston about 2.5 hours
Alternate route (our preference) would be to take I-93

north from Boston to I-495 north to Route 125 exit for
Haverhill, Mass/Plaistow, New Hampshire (be careful
as there are several 125 exits) Continue north on 125
to Rochester, New Hampshire and then north on US
16 to US 202 and signs for Sanford. After crossing
into Maine you will pass through Lebanon and then
up a long hill. At the top of the hill watch for Route
11a on the left. Turn left and it will take you down to
Springvale. Turn left again and proceed to Acton.
Driving time from Boston about 2+ hours.
183
The biggest difference is if you want services and

shops along the way. The I-95 route does not have
anything along the way whereas the 125 route,
although a bit slower traffic-wise, has a lot to see and
do, especially if you are looking for lunch!
To get to Acton from the Portland, Maine area take I-

95/Maine Turnpike south to the Route 109 exit for
Sanford and proceed as above.
Travel time from Portland is 1 hour.
Suggested Accommodations-
Nearest to Acton:
Motels
Mousam Valley Motel, Springvale, Maine 207-324-
2165
Campground
Apple Valley Campground 207-636-2285
In nearby Sanford, Maine, 9 miles south of Acton

Oakwood Inn & Motel, Sanford, Maine 207-324-
2160
Bar-H Motel, Sanford, Maine 207-324-4662
In South Sanford 15 miles from Acton
Super 8 Motel 207-324-8823
If you wish to stay at the ocean Wells, Ogunquit and

Kennebunk are all about an hour from Acton.
Kennebunkport is 1.5 hours from Acton (traffic can be
really heavy at times). There are literally hundreds of
places to stay along the coast
184
Recommended places to stay for Saturday evening in

Colebrook, New Hampshire
In downtown Colebrook:
Monadnock B&B - Bed and Breakfast
603-237-8216
Rooms With a View - Bed and Breakfast
603-237-5106
Colebrook House Motel & Restaurant
(603) 237-5521
Sportsmans Lodge and Cabins

Diamond Pond Road, Colebrook
(603) 237-5211
A bit remote and rustic but with great ambience and
food
Or for a real experience in old-time New Hampshire

luxury accommodations
The Balsam's in Dixville Notch
603-255-3000
Campground
Diamond Lake State Park Campground
There are many other campgrounds in the area.
185
NEW TAXA AND COMBINATIONS

Paul Martin Brown
A new combination is proposed here for the striking

white flowered form of the southern marsh orchid.
Although this species is restricted to northern
Newfoundland in North America, the white-flowered
form has been regular at the Cove area for many years.
The recognition of color forms at the forma level,
rather than the varietal level, is consistent with current
treatments of most genera.
Dactylorhiza praetermissa (Druce) Soo forma

albiflora (Druce) P.M. Brown comb. nov.
Basionym: Orchis praetermissa var. albiflora Druce,
R.B.E.C. 1919. p. 53
Synonym: Dactylorhiza majalis ssp. praetermissa
(Druce) D.M. Moore & Soó var. albiflora (Druce)
Harz
186
BOOK REVIEW
NATIVE ORCHIDS of NOVA SCOTIA
A Field Guide by Carl Munden
13 genera, 38 species including numerous varieties and forms
96 pages; 172 color photos; 19 line drawings; 43 maps
ISBN 0-920336-77-9 $39.95 CAD $27.00 USD + $4 postage &
handling
10% discount offered to NANOA members
order from: University College of Cape Breton Bookstore; PO
Box 5300,1250 Grand Lake Rd., Sydney, Nova Scotia, B1P 6L2,
Canada
For several years now Carl Munden has be

painstakingly, and not always so patiently, awaiting the
final product of his years of research on the native
orchids of Nova Scotia. The trials and perils of trying to
get a small work published, with color, are well known
to many, but finally Carl's book is available for all.
This 96-page (plus 44 pages of color) book

presents the orchids of Nova Scotia in a very readable
and easy to use format. Each species, and many
varieties, are presented with an excellent map showing
county distribution. Carl's notes on each species are
helpful both for identification and for the natural
history of the species. He makes not pretense for this
being a highly technical or taxonomic work, but despite
a few nomenclatural and taxonomic irregularities, it will
187
certainly serve as the best information available on the

family in Nova Scotia. His 172 color photos and
occasional line drawings help to illustrate most species
and varieties and will certainly help in their
identification. Following a recent trend in field guides,
no key is presented, although Carl has arranged the
species treatments by habitat and some natural
groupings. A glossary, brief bibliography and a narrative
chapter on hunting for orchids in Nova Scotia are also
included.
I would encourage all native orchid enthusiasts to

purchase the book in a timely manner, as the press run
is limited and I am sure they will soon be unavailable.
PMB
188
LOOKING FORWARD
September 2001
A New Species of Spiranthes from the

Southeastern United States
A Meeting with Ladies on Memorial

Day Weekend
Orchids, Whales, and Polar Bears
and more!
189

June 2001 North American Native Orchid Journal

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

Requests for either print or electronic copies should be

The 2001 orchid season is full upon us

There is no color in this issue as none of the

We will be in Maine through early October.

Paul Martin Brown

REPRODUCTIVE BIOLOGY OF THE

So different are lady's-slippers from other

This lip plays an important role in pollination.

Figure 1. Sketches of the flower of Cypripedium acaule. A. Side view

Although bees have been reported to sometimes

Daumann (1968) found that bees are well able to

Nilsson (1981) believes that the hairs inside the

below). Because these hairs are strongly light refractive,

In its escape the bee must pass two points where

The same bees are often trapped more than once

Botanists are uncertain just why bees enter the

that the bees feed on the hairs in the labellum (Nilsson

Stoutamire (1971) believes that bees must learn

However, Nilsson (1979) contends that, in

acetates and farnesene. The acetates are found in

A survey of 8 species of North American orchids

pollinators (Tengo 1979). A varying blend of odor

Among species of Cypripedium the size of the

The largest exit holes are found in the pink

labellum is produced on a short scape from a pair of

In keeping with the size of its exit holes, this

The establishment of bumblebee queens as

other Cypripedium is primarily dependent on bumblebee

A study in Ohio supports the role of Bombus and

Unlike the pollen vectors of most other lady's-

pollinators are present, and development of seed in the

Fruiting success in Cypripedium and other orchids

Many authors consider pollinator limitation to be

Williams 1966, Charnov 1982, Lloyd 1980, Waser and

Data based on hand pollinations suggest that an

on subsequent growth and reproduction (e.g., Lloyd

In Cypripedium acaule resource limitation resulted

Calvo and Horvitz (1990) also consider the costs

consistent with these predictions (Calvo 1993). A

Kull (1998) has reported observations on

slipper, C. passerinum among a mosaic of seres on the

According to Primack and Hall (1990) and

Artz, T. 1954. Frauenschuh und Spinne. Natur. Volk 84: 421-425.

Catling, P. M. and V. R. Catling. 1991. A synopsis of breeding

Faegri, K. and L. van der Pijl. 1971. The principals of pollination

Lloyd, D. G. 1980. Sexual strategies in plants I. An hypothesis of

Primack. R. B. and P. Hall. 1990. Costs of reproduction in the

Wright, L. W. 1975. The pollination ecology of Cypripedium acaule

Charles L. Argue, Department of Plant Biology, College of

Since press time for Native Orchids of the Southern

Until August 2000, this new species was known

On August 19, 2000, I returned to a site in Giles

four plants of Bentley's coralroot. This discovery

On August 26, 2000, friends from Pittsburg,

Searching later in September 2000, I discovered

At present then, there are six known Virginia

exceeded two hundred individual plants. Adding to the

Stan Bentley, 1201 MacGill St. Pulaski, VA 24301

BOOKS AND PAMPHLETS ON THE

The search for native orchids is a worthy goal,

Almost from the time I started studying wild