September 1998 North American Native Orchid Journal

NORTH AMERICAN NATIVE
ORCHID JOURNAL
____________________________________
Volume 4 September
Number 3 1998
a quarterly devoted to the orchids of North America
published by the
NORTH AMERICAN
NATIVE ORCHID ALLIANCE
* * * * * * *
* * * * * * *
IN THIS ISSUE:
PROCEEDINGS OF THE 3RD ANNUAL NORTH
AMERICAN NATIVE ORCHID CONFERENCE, 8-11
JULY, 1998, LAKE ITASCA, MINNESOTA - Part 1
WHAT MAKES A GOOD ORCHIDIST?……..and more!
ORCHID JOURNAL
(ISSN 1084-7332)
published quarterly in
March June September December
by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE,
Inc.
a group dedicated to the conservation and promotion of our
native orchids
Editor: Paul Martin Brown

Assistant Editor: Nathaniel E. Conard
Editorial Consultants:
Philip E. Keenan
Stan Folsom
Production Assistant:
Nancy A. Webb
The Journal welcomes articles, of any length, of both a scientific

and general interest nature relating to the orchids of North
America. Scientific articles should conform to guidelines such as
those in Lindleyana or Rhodora. General interest articles and notes
may be more informal. Authors may include line drawings,
and/or black and white photographs. Color inserts may be
arranged. Please send all inquiries or material for publication to
the Editor at PO Box 772121, Ocala, FL 34477-2121 (mid June -
August: PO Box 759, Acton, ME 04001-0759).
1999 Membership in the North American Native Orchid Alliance,

which includes a subscription to the Journal, is $26 per year for
United States addresses, $29US in Canada and $32US other
foreign countries. Payment should be sent to Nancy A. Webb, 84
Etna St. Brighton, MA 02135-2830 USA. Claims for lost issues or
cancelled memberships should be made within 30 days.
ORCHID JOURNAL
Volume 4 September
Number 3 1998
CONTENTS
NOTES FROM THE EDITOR
191
WHAT MAKES A GOOD ORCHIDIST?
The Slow Empiricist
193
PROCEEDINGS OF THE 3RD ANNUAL NORTH
AMERICAN NATIVE ORCHID CONFERENCE
8-11 JULY, 1998 LAKE ITASCA, MINNESOTA
Part 1
199
PROPAGATING NORTH AMERICAN CYPRIPEDIUM
SPECIES FROM SEED:
SUCCESSES AND PROBLEMS
Bill Steele
200
MANITOBA NATIVE ORCHIDS
Lorne Heshka
219
ALLOZYME VARIABILITY IN THE PLATANTHERA
CILIARIS COMPLEX
Nancy Cowden
241
ORCHID CONSERVATION IN THE 21ST CENTURY:
THE VALUE OF INCLUDING MYCORRHIZAL FUNGI
TO PRESERVE ENDANGERED SPECIES
Lawrence W. Zettler
261
PHOTOGRAPHING NATIVE ORCHIDS
Dianne Plunkett Latham
273
3RD ANNUAL NORTH AMERICAN NATIVE ORCHID
CONFERENCE FIELD TRIPS, JULY 10 & 11, 1998
Philip E. Keenan
285
LOOKING FORWARD:
December 1998
289
th
4 ANNUAL NORTH AMERICAN NATIVE ORCHID
CONFERENCE
290
Book Reviews
The Genus Cypripedium:
Natural History & Cultivation
Dr. Guido Braem, Charles & Margaret Baker
Wild Orchids Across North America
Philip E. Keenan
292
Unless otherwise credited, all drawings in this issue are by Stan Folsom
Color Plates:
1. p. 217 Cypripedium acaule, C. arietinum
2. p. 218 Cypripedium parviflorum var. pubescens flat-petalled type,
C. candidum
3. p. 259 Platanthera ciliaris, P. blephariglottis, P. xbicolor
4. p. 260 Spiranthes cernua seedlings
5. p. 271 Encyclia tampensis
6. p. 272 Corallorhiza maculata var. occidentalis forma immaculata,
Cypripedium reginae
7. p. 284 Platanthera praeclara, Malaxis paludosa, M. brachypoda,
M. unifolia
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 4, number 3, pages 191- 296; issued September 20, 1998.
Copyright 1998 by the North American Native Orchid Alliance, Inc.
Cover: Cypripedium fasciculatum by Stan Folsom
NOTES FROM THE EDITOR
This has certainly been quite a summer! With

raging fires in Florida, torrential rains in the Northeast,
extended heat in the Southwest and finally an August
when the weather has seemed to settle down and
proceed, but through it all there have been many
glorious native orchids. For 86 members of the Alliance
the 3rd Annual North American Native Orchid Conference in
northern Minnesota was certainly a highlight of the
summer. Many thanks are in order, but especially to
Lorne Heshka for arranging the trip to southern
Manitoba; Rob Freeman, Mark Larocque, and Dianne
Plunkett for scouting at Itasca; Char Menzel, Nancy
Webb, Nancy Cowden and many others who pitched in
and helped during the registration and the meeting
portion of the conference. Our extended thanks go to
all of the staff of Itasca State Park for their help and
thoughtfulness during our stay.
Two recent deaths in the orchid world should be

brought to note: Roberta Case, wife of Fred Case of
Saginaw, Michigan passed away this spring. She will be
remembered most recently as the co-author, with Fred,
of a great new book on Trilliums. Marion Ruff Sheehan
of Gainesville, Florida, one of the premier orchid
191
illustrators in the world, died in late July. Her work, with
that of her husband, Tom Sheehan, has been seen
monthly in Orchids (American Orchid Society Bulletin)
for many years as Orchid Genera Illustrated and has been
assembled recently in a book of the same name. That
was only a small portion of the work Marion did, not all
of it confined to orchids.
Plans are well underway for the 4th North American

Native Orchid Conference to be held next April in Florida.
Be sure to see the notice in this issue, and register and
make appropriate reservations early, as April is a busy
time in Florida.
Renewals will be sent out in mid September for

1999, with no increase in the membership/subscription
rate. Your early renewal helps us plan for the following
year. Please note that renewals will be sent to the office
in Ocala, Florida and all new memberships, claims and
orders for back issues to Nancy Webb in Brighton,
Massachusetts.
Paul Martin Brown,

editor
PO Box 772121
Ocala, Florida 34477-2121
tel/fax: 352/861-2565
email: naorchid@aol.com
192
Empiricist: WHAT MAKES A GOOD ORCHIDIST?
WHAT MAKES A GOOD ORCHIDIST?
The Slow Empiricist
At the recent 1998 NANOA Conference in

Minnesota at Lake Itasca, Paul Catling received the 1998
NANOA Education Award for his contribution to the
advancement of knowledge about native orchids.
Charles Sheviak and Carlyle Luer were similarly honored
at past conferences. These men have been shining lights
for the North American native orchid world.
These awards got me to thinking. Orchid

enthusiasts come in a variety of shapes, sizes and
abilities. What is important to one may not be of the
slightest interest to another. I contend that there is room
for each and every well-intended enthusiast. You will
notice that I qualified the enthusiast as being well
intended. I did this because there are those people who
are self-serving and will employ less than desirable
means to achieve their selfish goals. I am thinking of
those people who think nothing of digging up wild
populations to enhance their personal gardens or enrich
their herbaria. There are those who refuse to share
information out of professional jealousy. On a lower
level, there are those who usurp their field mates' time
with choice specimens by taking endless time with their
193
photography. There are other poorly-intentioned

behaviors but I would like to dwell on the positive
contributions the truly well-intentioned enthusiast brings
to the mix.
I have been impressed with many people I have

come in contact with since I started looking at orchids in
a serious way. I wanted to profile them in this column
but they have been for the most part very self-effacing
and shy about having themselves spotlighted. So instead
of causing them to be embarrassed by naming them
specifically, I have decided to highlight some of their
more desirable traits, traits that many good orchidists
share. I hope the models of these behaviors recognize
themselves and feel a sense of pride in that recognition.
As I first stated, enthusiasts come in many forms

and have many different motivations. There are the
serious students who are advancing our knowledge about
orchids with their studies. They work intensely with
them and have been exploring such avenues as gene
pools, chromosome counts and other very technical
aspects to determine new species and validate existing
ones. Their enthusiasm has increased our knowledge
extensively in the last few years. They have the
persistence and the patience to spend long hours
analyzing data that they have painstakingly assembled in
an effort to unravel some of the origins of certain
species. I salute you and applaud your efforts.
Then there are the people who are interested in

propagating the orchids and getting them to be
194
successfully grown in cultivation. They also have to have

perseverance, like the analysts of orchid structure, as they
struggle to understand the intricacies of the growing
medium, the needs of the emerging plants and all the
hairy problems that arise between the inception and the
fruition of their projects. You, too, are doing remarkable
things to advance our understanding.
Since my column usually champions the amateur

orchidist, I must discuss some of the outstanding
beginners I have encountered. Early in my own
excursions in the field, I came in contact with a woman
who was working to become more knowledgeable about
the plants she was seeing. Her method of learning was to
take a small notebook with her and perch beside the
plants and make a sketch of each new discovery. She did
not profess to be an artist but her skill was evident and
her detailed drawing gave her an intimate
acquaintanceship with her subject. What a wonderful way
to become more thoroughly cognizant of an orchid, or,
for that matter, any plant's uniqueness.
When I was a rank novice, I was also exposed to

much kindness on the part of those who had more
knowledge of the subject than I had at that time. One
woman, a botany professor at a Connecticut college,
approached her subject with a common-sense attitude
and down-to-earth explanations. She peppered her
narration with folksy good humor and was very flexible in
her acceptance of others. She had an appetite for
anything, and showed an interest in all things that would
195
come up. She is truly a light in my advance in

understanding.
Another woman who showed great sense gave me

the ability to accept what comes along with an up-beat
attitude. She had some mobility problems but she was
content to find things of interest in those areas that she
could reach, rather than bemoaning the fact that others
were farther afield. I have spent several excursions with
her when we elected to remain back in easier terrain. We
may have missed some specialties but we also had a
wonderful time exploring our own domain and often
found some things that the others missed.
Lest you think that all amateurs who have

anything positive to contribute are of the female gender,
I have had male field mates who have taken the time to
make sure that I saw a particular plant, or have slowed
their pace through the forest to accommodate my speed.
The more advanced males in knowledge about orchids
were often willing to take the time to explain an obscure
point that I might have missed. Some have shared their
loupes, their insect repellant, and their water when I was
in need.
To go back to the professional who contributes to

the enthusiasm about orchids, I would point out the
great teachers who are out there toiling to spread their
knowledge to others. One, in particular, was able to
make things clear in his explanations. His classroom
teaching style encompassed all his students' levels of
understanding and he made sure things were
196
comprehended before moving to a new area of

explanation. His field trips were always happy events
where each new discovery was enthusiastically examined
and enjoyed. He made everyone feel important and no
one was ever made to feel inadequate because of his
level of understanding.
Then there are the places of learning that

contribute to the empowerment of the enthusiast. Most
of the places I have encountered are interested in me
and try to provide me with the help I might need.
Whether it is as simple as a field station at a national
park dispensing information or a university with its
myriad tools like libraries, herbaria, etc., the people who
run these places usually try to give good, sound advice
and help to me in my quests. Their helpful attitudes are
sincerely appreciated when one is new to an area or
needs help of some kind.
Lastly, there are those who write about orchids.

They give of their knowledge and expertise to all of us.
This is a wonderful sharing of information. Whether you
are just an average enthusiast or a college-educated,
serious writer, you, through your words, have piqued
curiosity, enlarged comprehension, and added to the
enjoyment that shared experience brings to those who
are similarly engaged. You are a most important link
because you keep the flame alive and get the information
out to us.
To sum up, I would describe the good orchidist as

being one who is willing to share, who is deeply
197
committed to the field, and who wants to see the world

of information about orchids promulgated and
proliferated to all who are interested. I thank those who
have been positive influences for us. You know who you
are.
The Slow Empiricist
198
PROCEEDINGS OF THE 3RD ANNUAL

NORTH AMERICAN NATIVE ORCHID
CONFERENCE
8-11 JULY, 1998 LAKE ITASCA,
MINNESOTA
Part 1
PROPAGATING NORTH AMERICAN CYPRIPEDIUM
SPECIES FROM SEED: SUCCESSES AND PROBLEMS
Bill Steele
Lorne Heshka
ALLOZYME VARIABILITY IN THE PLATANTHERA
CILIARIS COMPLEX
Nancy Cowden
ORCHID CONSERVATION IN THE 21ST CENTURY:
THE VALUE OF INCLUDING MYCORRHIZAL FUNGI
TO PRESERVE ENDANGERED SPECIES
Lawrence W. Zettler
Part 2 in the December issue is scheduled to include:
Platanthera praeclara, a threatened prairie orchid
Margaret From
Recent Research on Minnesota Orchids
Welby Smith
Color, Form and Variation
Paul Martin Brown
Cypripedium hybrids of Mahnomen County, Minnesota
Rob Freeman
Piperias Unraveled
Mark Larocque & Paul Martin Brown
Recent Advances in the Systematics and Ecology of North
American Orchids
Dr. Paul M. Catling
199
Steele: PROPAGATING NORTH AMERICAN CYPRIPEDIUM
PROPAGATING NORTH AMERICAN

CYPRIPEDIUM SPECIES FROM SEED:
SUCCESSES AND PROBLEMS
Bill Steele
ABSTRACT
Eight North American species of Cypripedium have

now been grown to flowering from seedlings produced
in vitro by asymbiotic culture: pink lady's-slipper, C.
acaule, ram's-head lady's-slipper, C. arietinum,
California lady's-slipper, C. californicum, small white
lady's-slipper, C. candidum, spotted lady's-slipper, C.
guttatum, ivory-lipped lady's-slipper, C. kentuckiense,
yellow lady's-slipper, C. parviflorum, and showy lady's-
slipper, C. reginae. Seedlings of clustered lady's-
slipper, C. fasciculatum, sparrow's egg lady's-slipper, C.
passerinum, and yellow spotted lady's-slipper, C.
yatabeanum, have been produced by asymbiotic methods
and have been established ex vitro, but have, to my
knowledge, not yet been grown to flowering size.
Seedlings of the Central American species Mexican
yellow lady's-slipper, C. irapeanum, and downy yellow
200
lady's-slipper, C. molle, have been grown in vitro, but

have not been established outside of the flask. Only
very minor germination of mountain lady's-slipper, C.
montanum, seed has been achieved in the lab, and
seedlings planted out have survived for but one or two
seasons.
INTRODUCTION
Because of their showy appearance, orchids of the

genus Cypripedium have been prized in horticulture. All
too frequently, plants in cultivation have been obtained
by collecting from wild populations, sometimes to the
extent that such populations are jeopardized. Artificial
propagation of large numbers of these plants from seed
would thus serve simultaneously as a source for
horticultural material and as a disincentive for digging
wild plants. For this reason, researchers in North
America and also Europe and Japan have experimented
with artificial propagation of these orchids. Details of
the laboratory methods currently used for Cypripedium
and other terrestrial orchid propagation may be found in
Allen (1996). Production of seedlings in the laboratory
does not fully constitute propagation, however. The
seedlings must then be grown on to the flowering stage
for completion of the orchid life cycle, and for some
species the most difficult part of propagation is
establishing the plants out of the flask. An excellent
summary of present knowledge of horticultural
201
requirements can be found in Cribb (1997). In the

present report, I summarize the present status of in vitro
propagation of the North American Cypripedium species.
METHODS
Orchid seeds may be germinated in vitro either

symbiotically or asymbiotically. In symbiotic
germination, the seeds are inoculated with a compatible
fungus that provides nutrients to the orchid embryo, just
as occurs in nature. The researcher attempts to adjust
the cultural medium and conditions such that the fungus
grows sufficiently to nurture its orchid partner but not
sufficiently that the fungus overwhelms and kills the
orchid. In asymbiotic germination, the orchid seeds are
distributed on a sterile culture medium that provides the
nutrients that a fungus would provide in nature or in
symbiotic culture. To date, most of the success in
Cypripedium propagation has been obtained using
asymbiotic methods. All of my own work has been in
asymbiotic culture.
One of the main difficulties in germination of

Cypripedium seeds is that those of many species are
dormant at maturity. The purpose of the dormancy in
nature is to prevent germination of the seed at an
inopportune time such as just before the onset of winter.
Thus to germinate Cypripedium seeds, the dormancy must
somehow be circumvented.
There are two main ways to deal with the
dormancy problem: 1) sow the seeds in culture before
202
they are dormant, and 2) remove or neutralize the

biochemical agents in the seeds causing the dormancy.
Sowing the seeds before they are dormant,

sometimes loosely called ―green-podding‖ by orchidists,
requires that the seed be harvested in the short time
window during which the embryos are growing
vigorously before the pod parent introduces dormancy
factors into the seeds. A major difficulty in using
immature seeds is the determination of when to harvest
a seed capsule, for the window for successful embryo
culture is only a few days long.
In using fully mature seeds, the seed capsule is

harvested as close as possible to the time it would split
or ―dehisce‖ naturally. Mature seed of most Cypripedium
species apparently contains substances that prevent the
germination of the seed even when presented with all the
mineral nutrients and sugar necessary for growth of the
embryo. There are three methods used in an attempt to
overcome the dormancy: 1) stratification, 2) oxidation by
chemical bleaching, and 3) including a germination-
stimulating plant growth regulator or ―hormone‖ in the
culture medium. In stratification, the orchid seed is
maintained at a temperature near freezing for several
months. At this temperature, the seed seems to break
down germination-inhibiting substances metabolically.
Stratification seems more effective in some species than
in others.
203
The oxidation of germination-inhibitors is carried

out by immersing the seed in a bleaching solution prior
to sowing. The bleaching has the dual effect of
destroying microorganisms or spores on the seed that
would contaminate the culture as well as promoting
germination. In practice, two or even all three of these
methods are applied to promote germination of a given
batch of seed. In one species, C. montanum, no
combination of these methods has yet produced
successful germination of the mature seed.
PRESENT STATUS OF PROPAGATION EFFORTS
Table 1 summarizes the current status of artificial

propagation of the North American species. In this
table, the species are grouped into three classes
according to the relative difficulty of propagation on the
basis of not only the ease or difficulty of germination
and in vitro culture but of establishing out of the flask as
well. In the table, asterisks indicate species that have
been raised to the flowering stage from seed germinated
in vitro.
COMMENTS ON INDIVIDUAL SPECIES
All the species in the group I consider ―easy‖ to

propagate have been grown to flowering size from
seedlings produced in the laboratory. The species in this
category are both relatively easy to germinate and easy to
grow outside the flask. To my knowledge, the first lab-
204
grown seedlings of Cypripedium californicum to flower were

those raised by Michael Fong (personal communication)
of San Jose, California. His four-year-old seedlings
produced several flowers. Cypripedium candidum has been
grown to flowering by Allan Anderson (personal
communication) of the University of Guelph and
required nine years to flower. Carson Whitlow (personal
communication) of Adel, Iowa has told me that C.
kentuckiense typically requires five years to flower after
planting-out, and consequently I feel lucky to have had
two seedlings flower at four years from the flask. I have
had C. parviflorum var. parviflorum flower after two
growing seasons in pots, and I had one C. parviflorum var.
pubescens flower the second spring after removal from the
flask. In my experience, the latter subspecies more
typically requires three, four, or more years. As far as I
know the first publication of a Cypripedium to be
flowered from seed germinated in vitro was by Werner
Frosch (1985) of Germany, who raised C. reginae to
flowering the third year after sowing. Although I also
have had a seedling flower two years after planting-out,
three or four years seems more typical for C. reginae.
205
TABLE 1. Status of Propagation from Seed
Species Relatively Easy to Propagate:

C. californicum*
C. candidum*
C. kentuckiense*
C. parviflorum*
C. reginae*
Species Difficult to Propagate:

C. acaule*
C. arietinum*
C. fasciculatum
C. guttatum*
C. passerinum
C. yatabeanum
Species Presently Impossible to Propagate:

[C. dickinsonianum]
[C. irapeanum]
[C. molle]
C. montanum
*species that have been raised to flowering from seed germinated in

asymbiotic culture.
[ ] = Central American species.
206
pink lady‘s-slipper
Cypripedium acaule
207
pink lady‘s-slipper
Cypripedium acaule
208
ivory-lipped lady’s-slipper
Cypripedium kentuckiense
209
210
211
212
The group of species that I consider ―difficult‖ to

propagate is so primarily because of problems in
growing the seedlings once they leave the flask. All of
these species germinate fairly easily, and the major
difficulty lies in keeping the seedlings growing, or even
alive, after planting-out. I was delighted this May to
have a Cypripedium acaule seedling flower for the first
time, six years after planting-out. I may have been the
first person to do this. Approximately a week later,
however, Scott Durkee of New Haven, Vermont phoned
to say that he had a C. acaule flowering from seed only
two years after planting-out. My joy in my achievement
was somewhat deflated by Scott‘s superior
accomplishment! I have found that C. arietinum seed
germinates very readily under the appropriate conditions,
but seedlings of this species seem very sensitive to
reflasking shock and very often die from being moved
from one medium to another. If they survive reflasking,
C. arietinum seedlings may grow very large in
consideration of the relatively small size of the adult
plant and may be even larger than C. reginae seedlings at
the time of removal from the flask. The greatest
difficulties in raising C. arietinum seedlings is that they
require cool growing conditions, a very well-drained
medium, and relatively dry growing conditions. If all
these demands are met, I have found the seedlings will
flower the very next season after planting-out although
an additional year or two may be necessary. Cypripedium
guttatum also insists on cool growing conditions after
planting-out, but Dr. Svante Malmgren (personal
communication) in Sweden has carried this species
through to flowering. The other species in the
213
―difficult‖ group, viz., C. fasciculatum, C. passerinum, and

C. yatabeanum have not been flowered from seed to my
knowledge, but they germinate readily and can be grown
in pots, although with difficulty, and I have little doubt
that each will eventually be raised to flowering.
Cypripedium passerinum is probably the most difficult of
these species to grow outside the flask and seems to
require cool temperatures, or the light cycle typical of the
high latitudes from which the plant comes, or both in
order to survive.
Among the species that I consider ―impossible‖ at

present, Cypripedium montanum presents extreme difficulty
in the germination of its seeds. Despite many
experiments by a number of people, only a very few
seedlings have been produced by germination of mature
C. montanum seed. Peter Wilson (personal
communication) in England has perhaps had the greatest
success, but he was only able to obtain a few tens of
seedlings in many experiments. His limited success
involved no unusual treatments or media, and he feels
that success depends on the population or the clone
from which the seed comes. Immature seed of C.
montanum is also difficult to germinate, and I have only
ever obtained a handful of seedlings in this manner.
Karen May (personal communication), of Bigfork,
Montana has worked extensively at germinating
immature seed of this species. She has had good success
but with seed capsules from only a few of the many
plants from which she collected capsules, and she feels
214
that ease of germination varies greatly from clone to

clone.
The other presently ―impossible‖ species are from

Central America. Seed of Cypripedium irapeanum and C.
molle has been germinated and seedlings grown to the
stage for planting-out, but so far no one has been able to
grow these seedlings out of the flask. To my knowledge,
no significant efforts have been made to grow C.
dickinsonianum in vitro.
THE FUTURE
As growers gain more experience with lab-

propagated material, the ―easy‖ species should become
quite routine, and even now more or less casual
gardeners are successfully growing some of these species
from lab-propagated seedlings. Before long, someone
should succeed in blooming the ―difficult‖ species that
have not yet flowered. As more experience is gained
with ―difficult‖ species outside the flask, some of these
will also become routine, particularly in climates not
much different from those in which the plants are native.
Transplanted specimens of the Central American species
are being grown in cultivation in Mexico, but all of the
successful seedling production of which I am aware has
been outside that country. What would help greatly with
these species is either for seedlings to be shipped to
orchidists working near the natural range of these plants
or for Central American orchidists to grow the seedlings
in vitro.
215
Among North American species, Cypripedium

montanum seems to present the greatest challenge. The
plant is not that rare in the wild, so the seed must
germinate well in nature. Over a period of several years,
I found new seedlings under plants from which I
collected seed that I was unable to germinate. Clearly
there is a trick to germinating this seed, but all efforts to
find the secret have so far been futile.
REFERENCES
Allen, C., ed. 1996. North American Native Terrestrial Orchid

Propagation and Production. North American Native
Terrestrial Orchid Conference, Germantown, Maryland.
Cribb, P. 1997. The Genus Cypripedium. Timber Press, Inc.,

Portland, Oregon.
Frosch, W. 1985. Asymbiotische Vermehrung von Cypripedium

reginae mit Blüten drei Jahre nach der Aussat. Die Orchidee.
36: 30-32.
Bill Steele, Ph.D. Spangle Creek Labs 2295 County Rd. 445, Bovey, MN
55709 scl@uslink.net
216
SPECIES
Heshka: FROM SEED:
MANITOBA SUCCESSES
NATIVE ORCHIDSAND PROBLEMS
left:
First C. arietinum to flower 1 year
after planting out. April 1966.
below:
My first C. acaule to bloom - - 6
years after planting out. 17 May
1998.
Photos by Bill Steele
217
Heshka: MANITOBA NATIVE ORCHIDS
Top: Cypripedium parviflorum var. pubescens flat-petalled variant

Botton: Cypripedium candidum
Lorne Heshka Manitoba
218
218

Lorne Heshka
A total of 37 indigenous orchid species are listed

for the Province of Manitoba by Manitoba
Environment. Four of these species are not discussed in
this paper as I have not observed or photographed them
in Manitoba. These are:
Listera borealis – northern twayblade
Listera auriculata – auricled twayblade
Malaxis brachypoda – white adder’s mouth
Malaxis paludosa – bog adder’s mouth
One other orchid that I have not observed is

Cypripedium passerinum – sparrow’s egg or Franklin’s
lady’s-slipper. Jim Roy has lent me his slides of this
species and thus it will be covered in this presentation.
The province of Manitoba is located in a

transition zone between the Prairies to the west and the
Canadian Shield to the east. The province extends from
the US border at the 49th parallel in the south to the
North West Territories and the Arctic Ocean at Hudson
Bay in the north.
The vegetation distribution map of the province
shows grasslands occupying the south central and south-
western portion of the province. This area is primarily
219
agricultural, and except for protected areas, most of the

natural orchid habitat has been disturbed. From the
south east, in a diagonal band across the province, is the
parkland region, primarily a deciduous or mixed
deciduous and coniferous forest. Beyond are the lakes
and forest of the northern coniferous forest, occupying
the largest area of Manitoba. The most northerly portion
is the northern transitional forest including the Hudson
Bay lowlands and a thin border of tundra along
Hudson‘s Bay.
The largest number of species of orchids is found

in the southeast, while the largest numbers of orchids
can be observed further north in the northern coniferous
forest.
An increasingly popular recreational activity is

orchid viewing and orchid photography. This has put
some stress on the orchid habitat; however, the benefit is
an increased awareness of a need for conservation of
habitat for our native orchids.
eastern fairy slipper - Calypso bulbosa var. americana.

Our earliest-blooming orchid, and arguably the
most beautiful, is the eastern fairy slipper. This
species, although not common, seems to appreciate
a coniferous forest habitat growing over a limestone
base. The northern Interlake area provides this
habitat and in late May or early June you would be
able to find a number of blooming plants.
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ram’s-head lady’s-slipper, Cypripedium arietinum

This diminutive species can be found in two
distinctly different habitats. In black spruce and
tamarack bogs, this plant is normally found either as
individuals or occasionally in small clumps, while in
drier, often sandy locations, the plants can form
significant clumps. The small size of the flower as well as
the colour combine to make this a rather inconspicuous
plant in the dappled shade of its environment. One of
my favourite spots for this orchid is located in
Manitoba‘s Interlake region. Here the plants form nice
specimen clumps in open areas of a mature white spruce
grove.
Several orchids make their home in a Jackpine

Forest. These forests may be found on a sand base or
on the granite outcrops of the Canadian Shield.
pink lady’s-slipper - Cypripedium acaule

A common orchid in this habitat is the pink lady’s-
slipper, also known as the moccasin flower. This
widespread Cypripedium ranges from the southeastern
portion of the province to the Northwest Territories.
Most often found in dry pine forests or on the duff
covering granite of the Canadian Shield, it is occasionally
found in a black spruce and tamarack bog growing in
sphagnum moss. The plants in the bogs have always
been much reduced in size in comparison to those in
drier locations.
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ram’s-head lady’s-slipper,
Cypripedium arietinum
222
dragon’s mouth
Arethusa bulbosa
223
Approximately 15 years ago while canoeing with a friend

in the Nopiming Provincial Forest in eastern Manitoba
we came across a population of Cypripedium acaule that
included a scattering of the albiflorum or white-flowered
form. Fortunately I insisted we stop and record this on
film.
Hooker’s orchis - Platanthera hookeri

Blooming at the same time as Cypripedium acaule is
Platanthera hookeri. Although the green flowers of this
species are difficult to see, the two large leaves flat on
the ground may help. This plant seems to be equally at
home in a black spruce and tamarack forest where the
non-blooming plants may be confused with the pad
leaved orchis, Platanthera orbiculata.
northern slender ladies’-tresses - Spiranthes lacera

var. lacera
In July, after the Cypripedium acaule have long finished
blooming, the delicate spikes of Spiranthes lacera var. lacera
make their appearance. A good way to locate these
flowers is to crouch down and scan the forest floor for
the slender white spikes.
checkered rattlesnake orchis - Goodyera tesselata

At the same time and in the same areas you may find
Goodyera tesselata. Abundant when found in suitable
habitat, this, our largest Goodyera, colonises jackpine
forests on sand. In moist shady locations, plants up to a
foot high may be found.
224
small round-leaved orchis

Amerorchis rotundifolia
225
A common habitat through most of the North is the

black spruce and tamarack forest with underlying
base of sphagnum. This is a suitable habitat for a
number of orchid species.
lesser rattlesnake orchis - Goodyera repens

In a moist, often dark part of this forest, grows the
lesser rattlesnake orchis. This is the smaller of the
two Goodyera species we have and often occurs in small
colonies. The leaves of this plant, although rather small,
are the most attractive of any of our native orchid
species.
heart-leaved twayblade - Listera cordata var. cordata

Normally you could expect to find this species in
clean, mossy hummocks under a fairly dense black
spruce forest canopy. At my favorite site for this plant it
has as its companions ram’s-head lady’s-slipper, the
small round-leaved orchis and the blunt-leaved
orchid.
blunt-leaved rein orchis - Platanthera obtusata

The range of this small orchid in Manitoba extends at
least to Churchill on Hudson‘s Bay of the Arctic Ocean.
It is commonly found in black spruce and tamarack
forests throughout the province. I have observed the
foliosa form in at least three locations with a colony of
several plants in one particular location in southern
Manitoba.
pad-leaved orchis - Platanthera orbiculata
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The white flowers of this species stand out sharply in

the dimly lit shadows of the black spruce and tamarack
bogs where it is commonly found. I have photographed
this plant on Hecla Island on Lake Winnipeg in mid July,
and its range extends much further north.
green adder’s-mouth - Malaxis unifolia

This is the most common of the three species of
Malaxis in the province. Although common, this plant is
usually found widely scattered in an area. The flower is
best appreciated from the top allowing you to observe
each of the freshest flowers. The flowering stem on this
orchid elongates as it blooms with seedpods forming at
the bottom while there are still unopened buds at the
top.
small round-leaved orchis - Amerorchis rotundifolia

One of the earliest orchid species to bloom in
this habitat is small round-leaved orchis. Forming
dense colonies in favourable, usually quite moist,
locations within a black spruce forest, this dainty
orchid must be examined carefully to appreciate its
beauty. My wife tells me that when a magnifying
glass is used the individual flowers remind her of
angels adorned in white gowns spotted with pink.
This orchid is commonly found from the south
through to the extreme northern reaches of our
province.
Several of our most showy orchids live in a black

spruce, tamarack and sphagnum forest that has an
227
open canopy allowing good light to reach the

understory. One of the earliest showy orchids that
bloom in this habitat is the
dragon’s mouth Arethusa bulbosa

The vibrant color of this orchid makes it easy to find.
Never abundant in Manitoba, finding a group of three or
more is a rare treat. This orchid often competes for
attention with the grass-pink. The range within
Manitoba extends only as far north and west as the
northern points of our two largest lakes.
grass-pink - Calopogon tuberosus var. tuberosus

Years ago, before I caught the native orchid ―virus,‖
I found some pink petals on the ground in an open
sphagnum bog while photographing wildflowers. I
marked the spot, recorded the date, and made a point of
going back to this area the next year to check this out.
That was my introduction to the grass-pink and as you
can well imagine this new find fascinated me. Since that
time I have found colonies of this beautiful orchid in a
number of locations, the furthest north being at Hecla
Island although I understand their range extends even
further north than that. This year has been an especially
good year for grass-pink in our area and large colonies
exist where previously only single plants were found.
The Arctic tundra near the Port of Churchill on

the Hudson‘s Bay has only a few orchid species. The
most sought after species is the
228
229
sparrow’s egg or Franklin’s lady’s-slipper -

Cypripedium passerinum
This is our northern Cypripedium and although it is
probably found in locations south of Churchill, much of
its habitat is difficult to access. This orchid blooms in
early- to mid-July along with Platanthera obtusata and
Amerorchis rotundifolia.
One of my favorite orchid haunts is a fen

approximately 70 kilometers north of Winnipeg. Springs
of slightly alkaline water feed this fen and the open
meadows and surrounding forest provide habitat for
about twenty different species of orchids along with
other interesting plants such as butterworts, horned
bladderwort and pitcher plants.
large yellow lady’s-slipper - Cypripedium parviflorum

var. pubescens
Although numerous in this fen, it seldom colonises
an area in the same manner in which the small yellow
lady’s-slipper can. What it lacks in numbers, however, it
makes up in size. This was most dramatically illustrated
for me in June of 1996. My wife and I were introducing
friends of ours to the wonders of native orchids and had
stopped along a roadside where both the large and
northern small yellow lady’s-slippers were blooming.
We noted a particularly outstanding specimen on the
other side of a ditch filled with water. Once a route was
discovered across the ditch we were truly astounded by
the size of the blooms. The flowers on this plant
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measured 7.5" wide across the petal tips, 4" from top of
dorsal sepal to bottom of the pouch, and the pouch itself
was 2.25" long. All 5 flowers on this plant were of equal
size and the pouches were shaped more like a shoe than
a slipper having a flat bottom as opposed to the normal
rounded slipper shape. To provide a sharp contrast, a
northern small yellow lady’s-slipper was growing
immediately adjacent to this giant. Unfortunately when I
went to check on this plant this spring (1998) all I found
was a hole in the ground.
In particular environmental conditions, the petals

on this species are found to be flat, as opposed to having
the normal curl. My experience with the flat-petalled type
is limited to this fen where a mix of the normal flowers
were interspersed with a few of the flat-petalled types.
showy lady’s-slipper - Cypripedium reginae

This regal plant is the floral emblem of the province
of Prince Edward Island as well as for the state of
Minnesota. The showy lady’s-slipper appreciates the
sun for at least a portion of the day and in open
locations the plant is often quite short, while in heavily
shaded locations the plant will grow to nearly a metre.
This species requires an abundance of moisture and in
an optimum environment will develop significant
colonies. In 1996 I saw my first true albolabium, or white-
flowered form, of this species. These are truly impressive
plants and it is important that this colony be protected.
The colony is scattered over an area of approx. ½ km. by
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1 ½ km. including a black spruce bog as well as the fen.

Some stems have been observed with three blossoms.
hooded ladies’-tresses - Spiranthes romanzoffiana

This species has a wide range of habitat in
Manitoba. It is found growing on small islands in the
ponds of the fen as well as throughout the meadows of
the fen. It can be found in the sphagnum of the black
spruce and tamarack forest where the flower spike is
rather lax. In contrast, it grows in the tall grass prairie
where the flower spike is a compact spiral.
tall white northern bog orchis - Platanthera dilatata

var. dilatata
This tall white orchid is common in the open grassy
meadows of the fen. Companion orchids with a similar
blooming date in early July include Liparis loeselii and
Platanthera hyperborea as well as Platanthera huronensis.
northern green bog orchis - Platanthera hyperborea

var. hyperborea
This is a common to sometimes abundant orchid in
Manitoba in a variety of habitats with ample moisture.
Note: Paul Martin Brown advised that a photograph of a tall, stout plant of the
open fen was most likely Platanthera huronensis, the pale green bog orchid.
Loesel‘s twayblade - Liparis loeselii

Although this is a common orchid of the fen,
because of its size and color it is often difficult to find
among the grasses of the open meadows. It was quite by
accident that I first discovered this diminutive species.
Exploring a side drive off a gravel road in a boggy area,
232
three tiny plants caught my attention and we were able

to add another species to the list of 10 we had already
found in bloom that day. A careful check of the
surrounding area revealed a colony of some 40 plants.
In a wet floating bog in the southeastern region

of the province we find two of our less common
orchids.
rose pogonia - Pogonia ophioglossoides

At the extreme north-western extent of its range the
rose pogonia is not a common orchid in our province.
However, in appropriate habitat it is abundant and in
this particular bog which extends several kilometres, it is
impossible to estimate the number of plants except to
say that there must be many thousands.
In this same bog we find another very uncommon

orchid in Manitoba, the
green fringed orchis or ragged orchis - Platanthera

lacera.
Out in the open bog the flowers are a greenish
creamy white with deep fringes while the flowers on
plants found on the trail to this bog some 100 meters
away are green in color with shallow fringes.
Note: Paul Martin Brown suggested that some of the latter may be Platanthera
xandrewsii although the nearest known population of Platanthera psycodes is
approximately 150 kilometers away.
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A small colony of the plants along the trail were

associated with a larger colony of approximately 50
plants of Platanthera hyperborea. This species has been
recorded and positively identified as far north as Hecla
Island on Lake Winnipeg, approximately 300 km north
of the US border. The specific directions to this small
colony on Hecla were recorded. However, no one has
been able to locate these plants since they were first
observed some 20 years ago.
small purple fringed orchis - Platanthera pyscodes

Only one small colony of this species is known to
exist in Manitoba. This colony is located at the extreme
south east corner of the province and to reach this site
you are within 50 meters of the US-Canada border when
you leave the paved highway. Several years ago a
highway was built through the centre of the colony.
Fortunately a few plants survived in a moist deciduous
forest on either side of the highway.
northern bracted green orchis - Coeloglossum viride

var. viride
This orchid appears to prefer a dry habitat in a
deciduous or mixed deciduous/coniferous forest.
However this spring I found a couple of plants growing
within 20 centimeters of the edge of a small stream.
early coralroot - Corallorhiza trifida

This is the earliest of the saprophytic orchids to
bloom in Manitoba and is the most common. It also has
the most extensive range, from the US border through
234
to the northern border with the Northwest Territories.

The plants I have found have always been associated
with moist locations and I have even found them
growing in shallow water in a cedar bog. Whether they
actually grew in the water or were flooded after they
appeared could not be determined.
spotted coralroot - Corallorhiza maculata var. maculata

The bright red stems of this saprophytic orchid first
caught my attention in a small clearing in a mature white
spruce stand. All of the plants I have seen since have
been associated with white spruce. Commonly found in
cottage country, most cottage owners do not realize that
the flowers they are mowing are orchids.
striped coralroot - Corallorhiza striata var. striata

This member of the coralroot genus seems to be
much more tolerant of a variety of habitats than the
other Manitoba coralroots. I have found it in a pine
forest along with Cypripedium acaule, in mixed forests of
white spruce and aspen, as well as moist rich soils
adjacent to a small stream. The striped coralroot is
reasonably common throughout its range in Manitoba
and its striking appearance makes it easy to find.
northern small yellow lady’s-slipper – Cypripedium

parviflorum var. makasin
Readily forming large clumps, this orchid will quickly
colonise habitat it finds to its liking. In 1996 I stumbled
across just such a location near Lake Winnipeg,
approximately 70 km north of Winnipeg. While checking
235
out a location where I had, in previous years, found a

scattering of plants, to my astonishment, an east-facing
slope of a gravel road to a moist ditch contained literally
hundreds of plants. It appeared as though someone had
taken a pepper shaker and spread the seeds over a patch
approximately 5 metres wide by 12 metres long. A
follow-up visit to this site in 1997 found that little had
changed. This is one species of Cypripedium that can be
successfully rescued from a threatened area.
A single plant I rescued from a roadside before a

road grader‘s blade and transplanted to my yard, has,
within the space of a few years multiplied to over 90
blooming stems, many of which contain two flowers.
The fragrance emanating from this clump is
overwhelming.
The northern small yellow lady’s-slipper

normally appreciates relatively open sites. However, I
have found plants growing in fairly heavily wooded
locations. The plants in this type of habitat are often
single stems of considerable height, often growing in
excess of 20 inches.
small white lady’s-slipper – Cypripedium candidum

The flowers of this orchid tend to open early in
June while the leaves are still tightly curled about the
stem. An endangered species, this orchid unfortunately is
found in habitat that is most suitable for agriculture. It is
still found in several protected locations in Manitoba.
236
The Tall Grass Prairie is home to several colonies. A

companion orchid in the Tall Grass Prairie sites is
Great Plains ladies’-tresses – Spiranthes

magnicamporum
Blooming in mid-September, this plant is well hidden
among the Blue Stem Grass of the prairie. Its
wonderfully fragrant flowers do make it an easy orchid
to locate, as one can smell this orchid long before you
actually see it.
The final species is one of the orchids featured at

this conference, the
western prairie fringed orchis – Platanthera praeclara.

Although this species is found in only one area in
Manitoba, the numbers are incredible. In 1996 over
20,000 flowering plants were counted. The actual
number of plants in this area, including seedlings and
non-blooming plants, is difficult to comprehend.
Although the land where these orchids are found is
owned by a number of different organizations, the
overall management is overseen by the Nature
Conservancy of Canada.
Note of Interest:
Observation of Malaxis paludosa- the bog adder’s-
mouth in Manitoba
During the Friday, July 10, 1998 field trip to observe and
photograph the Malaxis paludosa near Itasca State Park, I remarked that an
area I frequently search for orchids, approximately 50 kilometres north of
Winnipeg was remarkably similar in many respects to the Itasca site. On July
237
13, 1998, Jim Roy of Winnipeg and I decided that we would check this area
for the bog adder’s-mouth. After 2 hours of unproductive searching we
decided to look in an ecological reserve on the opposite side of the highway.
We immediately found some nice specimens of Spiranthes romanzoffiana
and when I set up my tripod to photograph a plant, I noted three plants of
Malaxis paludosa directly below the camera. The “search image” from the
Itasca observations assisted in focusing on these plants.
The companion orchids within a metre of the Malaxis paludosa
were; Cypripedium arietinum – ram’s-head lady’s-slipper, Platanthera
hyperborea – northern green bog orchis and Spiranthes romanzoffiana –
hooded ladies’-tresses.
In discussion with Dr. Karen Johnson, the Curator of Botany at the
Museum of Man and Nature in Winnipeg, I learned that this species is
considered rare in Manitoba and there have been only 3 “confirmed”
observations in Manitoba, the last in 1985. There have been 7 other
“unconfirmed” observations. None of the previous observations have been in
this particular ecological reserve and none as near to the city of Winnipeg.
Lorne Heshka, 1204 De Graff Place,

Winnepeg, Manitoba R2g 1y8 Canada
e-mail: lheshka@escape.ca
238
239
Cowden: ALLOZYME VARIABLILITY IN THE PLATANTHERA
CILIARIS COMPLEX
above:
Platanthera blephariglottis
white fringed orchid
right:
Platanthera ciliaris
orange fringed orchid
240
CILIARIS COMPLEX
ALLOZYME VARIABILITY IN THE

PLATANTHERA CILIARIS COMPLEX
Nancy Cowden
The genus Platanthera consists of approximately

200 species distributed throughout the north temperate
regions of the world (Luer, 1975). North American
Platanthera taxa are assembled into five sections, some
more easily circumscribed than others, as Luer (1975) so
eloquently stated the matter. Among these intrageneric
groups, section Blephariglottis may arguably be the most
easily defined, with members having a distinctly fringed
lip, more or less. Examples of this assemblage include:
white fringed orchid, Platanthera blephariglottis, for which
the section was named, with its white flowers and
fringed labellum; orange fringed orchid, P. ciliaris,
similar to P. blephariglottis but characteristically orange in
color; and monkey-face orchid, P. integrilabia, white like
P. blephariglottis but bearing an only slightly serrated lip;
and purple flowered taxa such as large purple fringed
orchid, P. grandiflora. Within section Blephariglottis, the P.
ciliaris complex displays a particularly intriguing group of
variations on a common theme: floral color white,
241
CILIARIS COMPLEX
orange, or some intermediate shade, lips highly fringed

to barely serrate, individual flowers small (approximately
1.0 cm across the lateral sepals) or comparatively large
(2.0 cm or more across the lateral sepals), all typically
bearing 40 or more flowers per inflorescence.
In his study James Folsom delimited the

Platanthera ciliaris complex as P. ciliaris, P. blephariglottis, P.
cristata, and P. integrilabia as well as Chapman's hybrid
fringed orchid, P. xchapmanii, Canby's hybrid fringed
orchid, P. xcanbyi, and bicolor hybrid fringed orchid,
P. xbicolor, all hybrid taxa (Folsom, 1979, 1984).
Folsom‘s primary investigation considered the
appropriate taxonomic status of P. chapmanii, using floral
morphology, pollination biology, and cytology to
evaluate whether Chapman’s orchid should be
recognized at the specific level. Because P. chapmanii is a
purported hybrid between P. ciliaris and P. cristata,
Folsom focused his attention on these taxa, excluding
the remaining members of the complex (Folsom, 1979).
Morphological comparisons among the taxa that he
examined led Folsom to propose that P. chapmanii
functions as an independent species; however,
morphology could not provide conclusive evidence of
parentage (Folsom, 1984).
Taken as a whole, the floral morphological

variation and gradation among and within the taxa
comprising the Platanthera ciliaris complex would, I
suspect, make a thorough morphometric survey a true
242
CILIARIS COMPLEX
exercise in frustration. I found while working with P.

ciliaris and P. blephariglottis, as well as their presumed
hybrid P. xbicolor, that the only floral characteristic
capable of separating them from one another in a
principle components analysis was floral color (Cowden,
1993). Contrasting somewhat with my results, Judith
Robertson‘s study of disjunct populations clearly
demonstrated ecotypic differences in the floral
morphology and pollinator complements of coastal plain
and montane populations of P. ciliaris (Robertson, 1987;
Robertson and Wyatt, 1990a, 1990b). Although allied by
visible similarities, the P. ciliaris complex‘s evolutionary
relationship might better be evaluated by non-
morphological methods.
Generally speaking, Platanthera taxa for which we

have chromosome counts prove to have a somatic
complement of 42 chromosomes. There are exceptions,
as Sheviak and Bracht (1998) have determined for some
members of the P. hyperborea complex. For members of
the P. ciliaris complex, diploid chromosome counts prove
to be 42 for P. ciliaris, P. cristata, P. blephariglottis, and P.
chapmanii (Love and Simon, 1968; Folsom, 1979;
Dressler, 1981; Tanaka and Kamemoto, 1984). I am
unaware of reported chromosome counts for P.
integrilabia, P. xbicolor, or P. xcanbyi, but it certainly seems
reasonable to suspect that these taxa would possess
similar chromosome counts.
While morphology and cytology seem to confirm

what we suspect about relationships within the
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CILIARIS COMPLEX
Platanthera ciliaris complex, neither of these methods

provide the precision necessary to answer the question:
to what degree are these taxa related? Put a slightly
different way, does the P. ciliaris complex, as defined by
Folsom (1979), represents a natural assemblage of
closely related plants? In order to quantitatively evaluate
these questions, I employed starch gel, protein
electrophoresis.
As a method, protein electrophoresis has been

widely used by population biologists for over thirty years
but has seen few applications to orchid taxa. The
technique requires that active proteins, specifically
enzymes, be liberated from individual organisms, usually
by grinding up tissue in an appropriate extraction buffer.
The extract is then introduced to a gel matrix, often
either potato starch or polyacrylamide, and individual
enzymes are separated by electrical charge and/or
molecular size by running an electrical current across the
gel and its complementary buffer system. Afterward,
horizontal slices of the gel can be assayed for particular
enzymes by supplying the appropriate enzyme
substrates, cofactors, and other necessary components as
well as a dye that interacts with the specific enzyme‘s
activities (Wendel and Weeden, 1989). Variable forms of
the enzyme, known as allozymes if coded for by a single
gene or isozymes if coded for by different genes, appear
as bands that migrate different distances through the gel
matrix. Because distinct enzyme forms are inherited in a
Mendelian fashion, much like flower color, these
markers may be used to decipher mating patterns within
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CILIARIS COMPLEX
populations, as well as gene flow among populations and

among closely related species (Hamrick, 1989; Hamrick
and Godt, 1990).
For my study, leaf tissue was obtained from a total

of 21 populations of Platanthera taxa, including P. ciliaris,
P. blephariglottis, P. xbicolor, P. cristata, P. chapmanii, and P.
integrilabia. I was unable to sample material from P.
xcanbyi. When population size permitted a minimum of
twenty individuals per population were sampled. No
more than one lower leaf was removed from each plant;
in many cases one-half of the basal-most leaf provided
sufficient tissue for analyses. Enzyme activity in leaf
material was maintained by cold storage until assays
could be carried out. In all, nine enzymes, including 12
purported loci, were resolved using three buffer systems
(Table 1).
Banding patterns resulting from selective staining

procedures were used to infer allozyme variability.
Allelic designations for each individual were determined
by comparing Rf values to an internal standard run on
each gel. Unambiguous confirmation of allelic
comparability within and between taxa was achieved by
electrophoresing hypothesized variants side by side on
the same gel. Resulting banding patterns conformed to
those detected for other diploid plant taxa both in terms
of number of loci expressed and quaternary structure
(Wendel and Weeden, 1989; Weeden and Wendel, 1989).
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CILIARIS COMPLEX
Determinations of population-level allelic

frequencies, percent polymorphic loci, and alleles per
locus were performed using the computer program
BIOSYS-1, release 1.7 (Swofford and Selander, 1989).
Species-level allele frequencies were calculated by
combining data from all populations of a taxon. Percent
polymorphic loci and alleles per locus for species were
calculated according to the method of Hamrick and
Godt (1990). Species-level phenograms were generated
using the unweighted pair-group method with arithmetic
averaging (UPGMA) developed by Sneath and Sokal
(1973). Nei's unbiased genetic identity, I, (Nei, 1978)
was calculated for each taxon and provided values
utilized by UPGMA.
A total of 446 individuals from six Platanthera taxa

were analyzed. Five members of the Platanthera ciliaris
complex (P. ciliaris, P. blephariglottis, P. xbicolor, P.
chapmanii, and P. cristata) consistently shared at least one
allele per locus. The frequency of shared alleles varied
both within and among the five taxa, indicating no
taxonomically diagnostic variations.
Intraspecific comparisons of mean unbiased

genetic-identity values ranged from 0.777 in Platanthera
integrilabia to 0.999 in P. xbicolor. Mean genetic identities
among interspecifics were much more variable, with P.
blephariglottis and P. xbicolor sharing a mean value of 0.964
while the majority of pairs exhibited lower genetic
identities (Table 2). As is depicted in the UPGMA
phenogram of Nei‘s genetic identities, high genetic
246
CILIARIS COMPLEX
similarity values occurred among P. ciliaris complex

members (Fig. 1). First to cluster were P. blephariglottis
and P. xbicolor at 0.989, closely followed by P. ciliaris and
P. chapmanii which joined together at 0.971. The two
pairs of taxa link at the 0.930 identity level, and P. cristata
joined the group at 0.797. Also included in the P. ciliaris
complex, P. integrilabia clusters with the previous five
taxa at a relatively low value of 0.465.
Genetic similarity among Platanthera ciliaris

complex members is further illustrated by examining the
mean frequency of private alleles (Table 3). No unique
alleles were detected in either P. ciliaris or P. xbicolor. A
review of allelic frequencies on a locus-by-locus basis
shows that, with the exception of allele ―c‖ at PGM-
1(phosphoglucomutase, locus 1), P. blephariglottis
populations surveyed resolved no unique alleles. With
the aforementioned deviation, all P. blephariglottis
allozymes were also common to P. ciliaris, P. xbicolor, and
P. chapmanii. Private alleles were more frequent in other
members of the Platanthera ciliaris complex, including P.
cristata, with a mean value of 0.500 unique alleles per
locus and P. integrilabia, at 0.600 (Table 3).
247
CILIARIS COMPLEX
In practical terms, what do these results suggest?

First and foremost, with the exception of Platanthera
integrilabia, members of the P. ciliaris complex display a
very high level of genetic identity. Surveys such as those
reported by Gottlieb (1981) and Crawford (1983)
revealed that genetic identity values lower than 0.80 are
typical of congeneric taxa. A corollary of this assertion
is that conspecific populations regularly have genetic
identity values ranging from 0.80 to 1.00 (Gottlieb, 1981;
Crawford, 1983). While I would not suggest that P.
ciliaris and most of its complex members are
conspecifics, allozyme data point even more strongly
than do morphological observations to the conclusion
that P. ciliaris, P. blephariglottis, P. xbicolor, and P. chapmanii
are recent derivatives of a common origin. Platanthera
cristata, sharing a genetic identity with the
aforementioned taxa of 0.797, is likely also to have a
recent, shared evolutionary history. In contrast, P.
integrilabia has a comparatively low genetic identity value
(0.465) with other members of the P. ciliaris complex,
perhaps suggesting a more distant relationship.
At present, no other studies have compared

allozyme diversities among members of an orchidaceous
complex. However, there have been several studies that
investigated intrageneric genetic variation in terrestrial
orchids. Scacchi, De Angelis, and Lanzara (1990) found
that comparisons of genetic identity values among 11
Orchis species ranged from 0.000 to 0.767 with a mean
congeneric identity of 0.194. Low identity values were
also obtained by the same researchers for comparisons
248
CILIARIS COMPLEX
among three Epipactis species (Scacchi, Lanzara, and De

Angelis, 1987). An average congeneric identity of 0.197
was reported by Case (1994) for comparisons among
five North American Cypripedium species; the greatest
genetic similarity, 0.794, was found for the paring of
yellow lady's-slippers, C. calceolus (C. parviflorum), and
small white lady's-slipper, C. candidum, while the
lowest value, 0.000, was obtained for showy lady's-
slipper, C. reginae and ram's-head lady's-slipper, C.
arietinum. For the Platanthera taxa examined in this study,
all, except P. integrilabia, share genetic identity values
higher than those previously reported for other orchid
congeners.
Allozyme data do support the grouping of

Platanthera ciliaris, P. blephariglottis, P. cristata, P. chapmanii
and P. xbicolor into an assemblage conveniently referred
to as the P. ciliaris complex. However, P. integrilabia
proves to be a genetically more distinct member of the
genus. Is P. integrilabia sufficiently similar to the other
taxa examined to be grouped in the section Blephariglottis,
as defined by Luer (1975)? At this point data are
equivocal, needing a more thorough allozyme
investigation of other taxa included in this group. It is
clear that our concepts of intrageneric relationships in
the genus Platanthera will continue to be fine-tuned as
further investigations progress. This, undoubtedly, will
also prove true of other genera in the Orchidaceae.
249
CILIARIS COMPLEX
Nancy Cowden, Ph.D., Assistant Professor of Biology, Department

of Biology, Randolph-Macon Women's College, 2500 Rivermont
Ave., Lynchburg, VA 24503
Ncowden@rmwc.edu
250
CILIARIS COMPLEX
Literature cited
Case, M.A. 1994. Extensive variation in the levels of genetic

diversity and degree of relatedness among five species of
Cypripedium (Orchidaceae). American Journal of Botany 81(2):
175-184.
Cowden, N.E. 1993. A biosystematic study of Platanthera ciliaris, P.

blephariglottis, and P. xbicolor, emphasizing the northern portion of
their range. Ph.D. thesis. Miami University, Oxford, Ohio.
Crawford, D.J. 1983. Phylogenetic and systematic inferences

from electrophoretic studies. In S.O. Tanksley and T.J.
Orton [eds.], Isozymes in plant genetics and breeding, part A,
257-287. Elsevier Science Publishers, Amsterdam,
Holland.
Dressler, R.L. 1981. The orchids: Natural history and classification.

Harvard University Press, Cambridge, Massachusetts.
Folsom, J.P. 1979. Investigations into the nature of the putative hybrid
Platanthera xchapmanii (Orchidaceae). M.A. thesis. Vanderbilt
University, Nashville, Tennessee.
Folsom, J.P. 1984. A reinterpretation of the status and relations

of the taxa in the Platanthera ciliaris complex. Orquedia 9:
321-345.
Gottlieb, L.D. 1981. Electrophoretic evidence and plant

systematics. Annals of the Missouri Botanical Garden 64: 161-
180.
251
CILIARIS COMPLEX
Hamrick, J.L. 1989. Isozymes and the analysis of genetic structure

in plant populations. In D.E. Soltis and P.S. Soltis [eds.],
Isozymes in plant biology, 87-105. Dioscorides Press,
Portland, Oregon.
Hamrick, J.L. and M.J.W. Godt. 1990. Allozyme diversity in

plants. In A.H.D. Brown, M.T. Clegg, A.L. Kahler, and
B.S. Weir [eds.], Plant population genetics, breeding, and genetic
resources, 43-63. Sinauer, Sunderland, Massachusetts.
Love, A. and W. Simon. 1968. Cytotaxonomical notes on some

American orchids. Southwestern Naturalist 13: 335-342.
Luer, C.A. 1975. Platanthera. In The native orchids of the United States
and Canada excluding Florida, 176-242. New York Botanical
Garden, Bronx
Nei, M. 1978. Estimation of average heterozygosity and genetic

distance from a small number of individuals. Genetics
89:583-590.
Robertson, J.L. 1987. Pollination and reproductive biology of Platanthera

ciliaris in disjunct populations in the southeastern United States.
M.S. thesis. University of Georgia, Athens, Georgia.
Robertson, J.L. and R. Wyatt. 1990a. Evidence for pollination

ecotypes in the yellow-fringed orchid, Platanthera ciliaris.
Evolution 44: 121-133.
Robertson, J.L. and R. Wyatt. 1990b. Reproductive biology of

the yellow-fringed orchid, Platanthera ciliaris. American
Journal of Botany 77: 388-398.
Scacchi, R., G. De Angelis, and P. Lanzara. 1990. Allozyme

variation among and within eleven Orchis species (fam.
Orchidaceae), with special reference to hybridizing
aptitude. Genetica 81: 143-150.
252
CILIARIS COMPLEX
Scacchi, R., P. Lanzara, and G. De Angelis. 1987. Study of

electrophoretic variability in Epipactis helleborine (L.) Crantz,
E. palustris (L.) Crantz and E. microphylla (Ehrh.) Swartz
(fam. Orchidaceae). Genetica 72: 217-224.
Sheviak, C. J. and M. Bracht. 1998. New chromosome number

determinations in Platanthera. North American Native Orchid
Journal 4(2): 168-172.
Sneath, P.H.A. and R.R. Sokal. 1973. Numerical taxonomy. W.H.

Freeman, San Francisco, California.
Soltis, D.E., C.H. Haufler, D. Darrow, and G. Gastony. 1983.

Starch gel electrophoresis of ferns: a compilation of
grinding buffers, gel and electrode buffers, and staining
schedules. American Fern Journal 73: 9-27.
Swofford, D.L. and R.B. Selander. 1989. BIOSYS-1. A computer

program for the analysis of allelic variation in population
genetics and biochemical systematics. Release 1.7. Illinois
Natural History Survey, Champaign, Illinois.
Tanaka, R. and H. Kamemoto. 1984. Chromosomes in orchids:

counts and numbers. In J. Arditti [ed.], Orchid biology:
reviews and perspectives, III, 323-410. Cornell University Press,
Ithaca, New York.
Weeden, J.F. and N.F. Wendel. 1989. Genetics of plant

isozymes. In D.E. Soltis and P.S. Soltis [eds.], Isozymes in
Plant Biology, 46-72. Dioscorides Press, Portland, Oregon.
Wendel, N.F. and J.F. Weeden. 1989. Visualization and

interpretation of plant isozymes. In D.E. Soltis and P.S.
Soltis [eds.], Isozymes in plant biology, 5-45. Dioscorides
Press, Portland, Oregon.
253
CILIARIS COMPLEX
Werth, C.I. 1985. Implementing an isozyme laboratory at a field

station. Virginia Journal of Science 36: 53-76.
254
CILIARIS COMPLEX
Table 1. Enzyme systems, enzyme abbreviations, and associated

electrophoretic buffer systems used for allozyme resolution. Numbered
systems, with the exception of 8H*, appear in Soltis et al. (1983); LiOH is as
in Werth (1985).
Enzyme Enzyme Buffer

abbreviation system
Glutamate dehydrogenase- (E.C. 1.1.1.118) GDH LiOH,7
Glucose-6-phosphate isomerase- (E.C. 5.3.1.9) GPI 9,11
Isocitrate dehydrogenase (E.C. 1.1.1.42) IDH 11
Leucyl amino peptidase (E.C. 3.4.11.1) LAP 7,9
Malate dehydrogenase (E.C. 1.1.1.37) MDH 9
Phosphogluconate dehydrogenase (E.C. 1.1.1.44) PGDH 9,11
Phosphoglucomutase (E.C. 5.4.2.2) PGM 11
Shikimic acid dehydrogenase (E.C. 1.1.1.25) SkDH 8H
Superoxide dismutase (E.C. 1.15.1.1) SOD LiOH
Triose phosphate isomerase (E.C. .3.1.1) TPI 8H
*System 8H is a modification of Soltis et al. (1983) developed by

R. James Hickey. Gel and electrode buffers are as follows:
Gel buffer, pH 7.6 Electrode buffer,
pH 8.0
Tris...........……………………3.996g
LiOH............….1.636g
Citric acid, monohydrate......1.15 g Boric
acid.......16.262g
LiOH...........………………….0.168g Dilute to 1 liter
Boric acid.....…………………1.855g
Dilute to 1 liter
255
CILIARIS COMPLEX
Table 2. Matrix of mean genetic similarity (Nei's unbiased identity) for intraspecific (along the diagonal)
and interspecific (below the diagonal) comparisons among all Platanthera taxa. Range of genetic variation
appears in parantheses. N = number of populations for each taxon.
N ciliaris blephariglottis xbicolor chapmanii cristata integrilabia
ciliaris 9 0.908
(0.736-1.000)
blephariglottis 5 0.891 0.940
(0.712-0.991) (0.888-0.992)
xbicolor 2 0.896 0.964 0.999
(0.793-0.980) (0.933-0.998) (0.999-0.999)
chapmanii 2 0.889 0.853 0.855 0.895
(0.723-0.998) (0.728-0.961) (0.815-0.890) (0.895-0.895)
cristata 1 0.789 0.741 0.771 0.819 *****
(0.649-0.902) (0.663-0.875) (0.770-0.772) (0.806-0.829) (*****-*****)
integrilabia 2 0.471 0.466 0.460 0.433 0.307 0.777
(0.394-0.544) (0.393-0.550) (0.448-0.477) (0.398-0.503) (0.254-0.360) (0.777-0.777)
256
CILIARIS COMPLEX
Figure 1. Phenogram of relationships among Platanthera ciliaris complex

members, constructed from Nei’s unbiased identity values (Nei, 1978).
257
CILIARIS COMPLEX
Table 3. Proportion of polymorphic loci (P) and mean number of

alleles per locus (A), for populations (p) and taxa (t) and the
average frequency of private alleles per locus (p(1)) for each taxon.
Platanthera taxon Pp Pt Ap At p(1)

ciliaris 0.305 0.666 1.324 1.833 0.000
blephariglottis 0.216 0.500 1.250 1.583 0.030
xbicolor 0.250 0.250 1.250 1.250 0.000
chapmanii 0.500 0.583 1.541 1.667 0.142
cristata 0.166 0.166 1.166 1.166 0.500
integrilabia 0.291 0.500 1.333 1.583 0.600
258
CILIARIS COMPLEX
259
Zettler: ORCHID CONSERVATION
260
ORCHID CONSERVATION IN THE 21ST

CENTURY: THE VALUE OF
INCLUDING MYCORRHIZAL FUNGI TO
PRESERVE ENDANGERED SPECIES
Lawrence W. Zettler
Introduction
Orchids have evolved a unique mycorrhizal

symbiosis in which the fungal partner is consumed as an
energy source by the orchid (=mycotrophy). In nature,
the orchid life cycle is completely dependent on the
availability of specific fungi to initiate seed germination
and seedling development. As orchid habitats continue
to be destroyed by human-related factors, a pressing
need arises to secure mycorrhizal fungi for use in orchid
conservation efforts projected for the next century. This
paper will briefly describe the orchid-fungal symbiosis,
explain how fungi can be used to grow orchids from
seed, and emphasize the importance of the symbiosis for
conservation during the 21st century.
The Orchid-Fungal Symbiosis and its Significance
The orchid-fungal symbiosis begins when an

orchid seed‘s embryo becomes infected by a Rhizoctonia-
like fungus (Basidiomycotina). Upon entering the
suspensor region of the embryo, the fungus proliferates
261
within cortical cells of the host, forming ball-like masses

of hyphae, termed pelotons (Fig. 1). Pelotons are
distinctive structures unique to an orchid mycorrhiza,
and are usually confined to cortical regions of root-like
organs. In a stable symbiosis, orchids are able to control
the spread of fungal infection by producing natural
fungicides (phytoalexins) and by digesting pelotons (Fig.
2). The latter process yields carbohydrates (energy),
previously tied up in fungal hyphae, to the host orchid.
As such, orchids are mycotrophs or ―fungus feeders.‖
Because there is no evidence that the fungus benefits
from this association and it appears to be harmed but
rarely killed, most authorities now view the symbiosis as
a form of parasitism by the orchid on the fungus. The
ability of orchids to obtain energy from fungi, in
addition to sunlight (photosynthesis), clearly
demonstrates that these plants are heterotrophic,
especially as seedlings. Once mature, orchids are
believed to exist as both heterotrophs and autotrophs,
depending on a given species‘ needs. With this
alternative nutritional capability, the Orchidaceae has
evolved to become the world‘s most diverse plant family
with an estimated 30,000+ species. Orchid seeds, for
example, have no need for weighty food reserves
(endosperm) making them capable of long-distance wind
dispersal into new habitats favoring speciation. As
another example, some terrestrials such as pink lady's-
slipper, Cypripedium acaule, perform ―disappearing acts,‖
i.e., they remain below ground for several years as
mycotrophs, then reappear in an autotrophic capacity
perhaps to maximize reproductive success in favorable
years.
262
While the utilization of fungi as an energy source

appears to have benefited orchids as a whole, it has its
costs. Without fungi in the natural habitat, seeds would
fail to germinate, abruptly ending the orchid life cycle.
This absolute requirement for fungi has made growing
orchids from seed a complicated and difficult process
compared to other plants. For endangered orchids, this
dilemma poses a serious challenge to conservation
efforts. Artificial attempts to propagate orchids from
seed on nutrient media without fungi have been largely
successful for the epiphytes (Fig. 3) but unsuccessful for
many terrestrials. If orchids are artificially propagated
with little regard for fungi and reintroduced into
―restored‖ habitats, they will not be able to spawn
seedlings unless suitable fungi are available for
germination. As native orchid habitats continue to be
destroyed by human-related factors, there arises an
urgent need to isolate, screen, and preserve mycorrhizal
fungi. One means to propagate orchids and screen fungi
for conservation purposes is to employ the technique of
symbiotic seed germination.
263
264
Symbiotic Seed Germination: An Overview
Symbiotic seed germination seeks to mimic nature

by exposing orchid seeds to fungi to initiate the
germination process, usually in the laboratory. It is
considered a valuable conservation tool because it can be
used to rapidly germinate seeds of endangered species
that often resist attempts at artificial (asymbiotic)
propagation. It also provides a means to screen fungi
for their effectiveness at initiating a symbiosis. In
addition, it permits fungal-infected seedlings to survive
soil transfer better than those grown without fungi. Its
primary drawback lies in the investigator, because few
are trained in mycology or are willing to work with two
organisms (orchids and fungi) instead of one (the
orchid).
The most widely used symbiotic technique

involves sowing surface-sterilized seeds in vitro over a
low-nutrient medium such as oatmeal agar within a Petri
dish or test tube slant. Upon the introduction of an
effective fungus followed by incubation in darkness,
orchid seeds often germinate rapidly (ca. 2 weeks) and at
high percentages (>50%), depending on the species and
seed pretreatment used. Under optimal conditions,
orchid embryos undergo rapid and dramatic change in
size and appearance, transforming into leaf-bearing
seedlings from masses of undifferentiated cells. Most
seedlings can be exposed to white light upon leaf
expansion until tissues develop green pigmentation and
presumably initiate photosynthesis. Eventually, such
seedlings are suitable for transfer ex vitro, and usually
265
survive provided they are acclimated in a greenhouse on

a mixture of moist, pre-inoculated soil and Sphagnum
moss (Fig. 4). In cooler climates, seedlings should be
transplanted into suitable habitats during spring, after
the threat of frost. Presumably, fungal-inoculated
seedlings have the ability to digest pelotons as a food
source, making their transition to adulthood more likely.
It is also assumed that fungal inoculum from the
seedlings may permeate into the surrounding substrate
and spawn additional seedlings.
In 1993, a new symbiotic technique was

developed that has the potential to revolutionize our
understanding of orchid ecology and greatly benefit
conservation efforts. It involves placing terrestrial
orchid seeds in packets constructed from nylon plankton
netting followed by burial in a natural substrate (Fig. 5).
Mycorrhizal fungi in the substrate are permitted to pass
through the netting and into the orchid embryos.
Packets are then retrieved and inspected for fungal-
infected seedlings. For the first time, fungi can be
isolated from young seedlings in the natural habitat—a
process that would practically be impossible considering
the minute size of seeds and seedlings. Clearly, this
technique provides a quantitative means to study orchid
seed germination in situ. It also represents an instrument
Zettler used to extract the most effective fungal isolates
responsible for germination. To a mycologist, the
technique can be viewed as a way to ―bait‖ fungi from
soil. Once extracted from orchid tissues, fungi can be
screened for their effectiveness at promoting in vitro
germination, followed by permanent deposition into an
266
international culture collection such as the University of

Alberta Microfungus Collection and Herbarium
(UAMH). In light of ongoing habitat destruction, I
strongly urge other investigators to consider using both
techniques in order to preserve the often-overlooked
fungal component.
Conclusion
The ecology and distribution of orchid

mycorrhizal fungi in nature remains poorly understood
despite their importance to the Orchidaceae. Recent
evidence suggests that orchid fungi may persist as
independent saprophytes with a wider distribution than
the orchids they support. Some orchid fungi may even
form partnerships with members of other plant families.
Until it can be demonstrated that habitat destruction
does not contribute to the loss of the fungal taxa
required by orchids, every attempt should be made to
include fungi in conservation efforts. This includes the
fungi of epiphytic and terrestrial taxa alike, despite the
relative ease by which the former group can be cultivated
from seed in the absence of fungi. It is simply too easy
to focus on the highly visible orchid plant while ignoring
the rarely seen, poorly understood, yet critical fungal
component. The use of in situ symbiotic seed
germination provides hope for isolating the most
effective fungi. In vitro symbiotic techniques provide the
method to screen fungal isolates, and cultivate orchid
seedlings with a mycotrophic capability.
267
Acknowledgments
I extend thanks to Marta R. Bowen (Springfield, IL) for critiquing the text, and
to Paul Martin Brown for providing the photograph depicted in Fig. 3.
FURTHER READING:
Arditti, J. 1966. Orchids. Scientific American 214: 70-78.
Clements, M. A. 1988. Orchid mycorrhizal associations. Lindleyana 3: 73-86.
Gill, D. E. 1996. The natural population ecology of temperate terrestrials: pink

lady‘s-slippers, Cypripedium acaule. In: North American native terrestrial
orchid propagation and production (conference proceedings), National
Arboretum, Washington, D.C. (ed. C. Allen) pp. 91-106.
Rasmussen, H. N. 1995. Terrestrial Orchids: From Seed to Mycotrophic Plant.

Cambridge University Press, Cambridge.
Rasmussen, H. N. and D. F. Whigham. 1993. Seed ecology of dust seeds in

situ: A new study technique and its application in terrestrial orchids.
Am. J. Botany 80: 1374-1378.
Rasmussen, H. N. and D. F. Whigham. 1998. Importance of woody debris in

seed germination of Tipularia discolor (Orchidaceae). Am. J. Botany 85:
829-834.
Zelmer, C. D. and R. S. Currah. 1995. Evidence for a fungal liaison between

Corallorhiza trifida (Orchidaceae) and Pinus contorta (Pinaceae). Can. J.
Botany 73: 862-866.
Zettler, L. W. 1997. Orchid-fungal symbiosis and its value in conservation.

McIlvainea 13: 40-45.
Zettler, L. W. 1997. Terrestrial orchid conservation by symbiotic seed

germination: Techniques and perspectives. Selbyana 18(2): 188-194.
Lawrence W. Zettler, Ph.D., Department of Biology, The Illinois College, 1101 West
College Ave., Jacksonville, Illinois 62650 USA. Dr. Zettler teaches organismal
biology at The Illinois College and directs undergraduate student research in
collaboration with The Morton Arboretum in Lisle, IL where he serves as a
Research Associate.
E-mail: lwzettle@hilltop.ic.edu
268
269
DON'T FORGET THE 1999
4TH NORTH AMERICAN NATIVE

ORCHID CONFERENCE!
LARRY ZETTLER WILL BE

SPEAKING AGAIN AND UPDATE US
ON CURRENT SUCCESSES WITH
SPECIFIC SPECIES OF OUR NATIVE
ORCHIDS
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270
Encyclia tampensis
Florida butterfly orchid
FL P.M. Brown
271
271
Latham: PHOTOGRAPHING NATIVE ORCHIDS
Left:
Corallorhiza maculata var.
occidentalis forma
immaculata
western spotted coralroot,
yellow spotless from
MN
Below:
Cypripedium reginae
showy lady's-slipper
MN
272
Photographing wildflowers such as native orchids

need not be difficult or involve a large amount of
expensive equipment. Whereas an avid nature
photographer could use a triple flash/tripod set-up with
a diffusion tent, I don‘t recommend it, as you may need
to bring along a Sherpa to tote all your equipment into
the field! To spare my back (and the environment) I use
a minimum of equipment. Depending on the film speed
preferred, a tripod may be unnecessary. I recommend
basic equipment as follows: a single lens reflex camera
(which is NOT set on automatic), a flash unit, and
magnification equipment such as a macro lens or
stacking close-up lenses. If you have a little extra space
in your backpack, I would also add one or two of the
commercially available expandable reflectors.
The Camera
A single lens reflex camera is necessary so that
you can preview your photographs before snapping the
shutter and confirm that the subject of interest is in
focus. Do not use the automatic setting if you happen
to have that feature, as camera light meters can be fooled
273
in the dim light of the forest floor. It is better to log

your shots for a few rolls. That is, keep track of your
shutter speeds and F stops for each photograph. Review
the photographs with the settings in various light
conditions, thereby learning what range of settings work
well for the film speed you prefer. You should then
bracket your shots from among the range of useful
settings. For example, if you are shooting in full sun,
and you want maximum depth of field, a standard setting
is achieved as follows. Set the shutter speed equal to the
ASA of the film you are using (i.e., ASA 64 is 1/60 of a
second) and use an F stop of 16. To bracket this you
could either vary the shutter speed setting (1/30 or
1/125 for example), or you could vary the depth of field
setting (F11 or F22 for example). Be careful if you vary
both the shutter speed and the F stop as you may cancel
out any change in lighting. A shutter speed of 1/125 at
F8 admits the same amount of light as a shutter speed of
1/ 60 at F11, for example. The only difference one
would see in these settings is that of a greater depth of
field at the F11 setting. The higher the F stop, the
deeper the area which is in focus. It is not unusual for
me to shoot a 24-exposure roll per blossom trying many
different settings. Somewhere on the roll you will
indeed get the perfect shot! An automatic camera will
not let you have the necessary level of control.
Flash
You will not need a very powerful flash as most
wildflower photographs are taken at close-up range. To
determine the correct setting for a flash exposure, you
274
could review the flash specifications, dust off your

physics formulas on the fall-off of light, and measure
your distance to the subject. You can then precisely
determine what shutter speed and F stop is correct for
your speed film at a specific distance. That would be too
much work for me, however. It is far easier to log a
range of settings on a roll of shots using your preferred
film and flash. From this you can tell what settings work
best at your typical working distance for wildflower
close-ups.
Some photographers set up two auxiliary flashes

mounted on tripods at either side of the subject. This is
in addition to the main flash on the camera, which is also
mounted on a tripod. A triple flash set-up will yield a
very balanced light. Auxiliary flashes are activated by a
slave unit attached to the auxiliary flash. The slave unit
senses the burst of the main flash and trips the auxiliary
flash. This necessitates three tripods (and a Sherpa)
however, to say nothing of disturbing the environment
while setting up all the tripods. I don‘t recommend this
type of tripod set-up in fragile areas such as where
orchids typically grow.
Close-up Equipment
To achieve sufficient magnification, a set of
stacking close-up lenses, a macro lens, a reversal ring, a
bellows, a set of extension tubes, or combination
thereof, is necessary for wildflower photography. The
least expensive way to achieve magnification is to use
275
close-up lenses. A typical set of three close-up lenses can

be purchased for less than $35.00. This is also one of
the lightest weight methods of achieving magnification.
A typical set includes magnification lenses of +1, +2
and +4. Such a set of close-up lenses can be combined
for +3, +5, +6, or +7 magnification. The least light is
lost with this method of magnification because the focal
length is not significantly extended. Most of my orchid
photographs were taken with these.
I presently use a 90mm Tamron macro lens. With

the increased focal length of the macro lens, the working
distance is greater from the subject to the camera flash
than with close-up lenses. As a consequence, the F-stop
must be opened wider when using a macro lens since the
camera with flash attachment will be back farther from
the subject.
A reversal ring is another inexpensive, light-

weight method of achieving magnification. With a
reversal ring, take the regular lens, in my case an F1.8,
55mm lens, reverse the direction of the lens, and attach
the lens by screwing it on to the barrel of the camera
using the reversal ring. This will yield about a one-to-one
life-size photograph. Reversing a 24mm wide-angle lens
will yield about a 2.5 magnification. This is the
magnification method used by Lorne Heshka for his
excellent program on Manitoba's Native Orchids during
the July 1998 NANOA conference. An advantage of
reversing the lens is that no additional light is necessary.
276
The disadvantage of a reversal ring is that you cannot

use the lens focusing system but must focus by moving
the camera, thus it is convenient to have a focusing rail.
With the reversal ring you lose the bright wide open lens
focusing because the automatic diaphragm lever is facing
away from the camera. You can use your finger,
however, to push this lever to open the lens diaphragm
for focusing and release it to stop-down mode when
releasing the shutter.
Another alternative for magnification is to use a

bellows and rack out the lens to the desired length for
the intended magnification. Using a bellows has the
advantage of allowing magnification on a sliding scale
rather than a fixed magnification system such as when
reversing a lens. A bellows has the disadvantage of
being heavy, bulky, time consuming to set up, and
requires more light because of the increased focal length.
A final magnification option to consider is to add

one or more extension tubes. This could be done in
combination with the bellows. This has the
disadvantage of increasing the focal length, thereby
necessitating greater illumination. Extension tubes are a
relatively lightweight, relatively inexpensive way of
achieving magnification.
Reflectors
277
If you do not want to use an auxiliary flash, it is

wise to add some reflectors to your backpack to bounce
in additional sunlight. Compact, expandable reflectors,
which fold into a small area, are commercially available.
When released, they form a circle of white, silver or gold
material. I use the LITEDISC 12 inch diameter white
reflector and the 22 inch diameter white/silver reflector
manufactured by Photoflex. They fold to a five-and-one-
half inch diameter and a 9-inch diameter size
respectively. Alternatively, one can make his or her own
reflector by covering any size or shape piece of
cardboard, or stiff material, with a foil such as Reynolds
Wrap. If you don‘t have a friend to hold the reflectors
for you, be sure to bring a few of your gardening sticks
to lean them against. Bamboo sticks take up little room
in your pack and can be driven into the ground with
minimum impact to the environment
.
Film
I have a strong preference for Kodachrome 25
film because it provides the greatest resolution (detail).
Such slow film, however, will generally require the use of
a tripod, unless the subject is in quite a bright light. In
general, you want to use the slowest film available that
you are willing to consider in order to obtain the greatest
detail. The slower the film, the greater the detail. If
Kodachrome 25 is too slow for your patience, try
Ektachrome 64. Higher speed film is useful if there is
much wind, or if conditions are very dark, and you do
not want to increase light artificially.
278
Tripod
A tripod is advisable, especially if you are using
slow-speed film in low-light conditions. I use a Gitzo
tripod with a Bogen head. The advantage of a Gitzo
tripod is that the legs can be spread out nearly flat to the
ground. The advantage of the Bogen head is that it has a
shorter column than the standard Gitzo head, allowing
the tripod to be positioned close to the ground. The
tripod head can be swiveled and tilted in many
directions. Using a strap to sling the tripod over your
shoulder will make hiking easier. When working with a
tripod, using a cable release is always recommended, to
eliminate any movement of the camera.
Natural Backgrounds
Having a complementary yet contrasting
background is the objective of wildflower photography.
This can be done by several means. Blurring the natural
green background is very pleasing. A pink lady’s-
slipper, Cypripedium acaule, photographed at St. Croix.
State Park in Minnesota with a +6 close-up lens was
taken with Kodachrome 25 at a 1 second F16 exposure
using a tripod and flash. The one-second exposure
brings the light from the green background through. An
alternative example showing more of the background in
another pink lady’s-slipper photograph taken at Land
O‘ Lakes State Forest in Minnesota using Kodachrome
25 with a one second exposure at F16 with a flash. The
279
detail is beautiful, but a very still day is required for these

exposures!
Black Backgrounds
An alternative to using a background that shows
the color of the environment, is to use settings which
produce a black background. Although I find it more
pleasing to use the color of the natural background, this
is not always possible. In the attached showy lady’s-
slipper, Cypripedium reginae, which I photographed at Ft.
Snelling State Park in Minnesota, I had a wire fence in
the background. To eliminate distracting backgrounds,
choose settings such that the fall-off of light will result in
a black background. Black backgrounds will also give
the maximum contrast for those photographs in which
you want the subject to stand out with maximum impact.
The following sequence shows the variety of equipment
and film that can be used to achieve a black background.
The C. reginae black background which eliminated the
fence was achieved through the use of Ektachrome 64
and an exposure of 1/60 of a second at F22 with a flash.
I call this type of a shot a ‗grab‘ shot because the camera
can be hand held without the need of a tripod. A +4
close-up filter was used as well. A similar black
background effect can be achieved by using
Kodachrome 25 with a 1/60 second F16 exposure using
close up filters as was done with a western prairie
fringed orchid, Platanthera praeclara. A black
background was also achieved in the attached western
spotted coralroot, yellow spotless form, Corallorhiza
maculata var. occidentalis forma immaculata, taken at Scenic
280
State Park in Minnesota, using Kodachrome 64 with a

one second, F16 flash exposure, including a 90mm
macro lens with 3 extension tubes plus a tripod.
Another example of a dark background was achieved
with a Calypso taken on Flower Pot Island near the Bruce
Peninsula in Lake Huron, Ontario. It was shot with
Kodachrome 25 at a ½ second, F22 exposure with flash
and a tripod using +6 close-up lenses. In summary,
when using Kodachrome 25 and close-up lenses, a one
second F16 exposure will bring through the background
whereas a 1/60 second F16 exposure at a similar
magnification and flash will cause the background to
black out.
Artificial Backgrounds
An example is of a Case’s ladies’- tresses,
Spiranthes casei, that showed the ditch habitat near
Spooner, Wisconsin. It was taken on Kodachrome 25
with a one second, F16 exposure using a tripod and
flash. To achieve a more effective close-up and remove
distractions in the background, I had a friend hold the
blue reverse side of my jacket in the background. The
photograph was taken at a one second, F11 exposure
with a +6 magnification on a tripod with a flash.
Another technique for replacing distracting backgrounds
is to move to your setting something natural from
elsewhere in the environment. Examples could include a
piece of birch bark or a moss covered rock. Be sure to
return whatever you have chosen to the spot from which
you found it so as to have minimal impact on the
environment. When photographing in the field, be
281
careful not to trample immature plants and other

companion plants. Resist the urge to snip off the
flowers of adjacent plants which may be a distraction in
your background; a native plant population needs the
flower‘s seeds to sustain itself. Instead, use one of the
artificial background or black background techniques,
which I have described, to remove any distracting
elements.
Composition
There are some important principles to keep in
mind when composing your photograph. These include
positioning of the center of interest, positioning leading
line diagonals from the left side of the photograph, and
selecting an odd number of subjects. Resist the urge to
place a subject in the center of a photograph, as such a
composition is static. It is better to place a subject on
one of the ‗golden thirds‘. For this placement, imagine a
tic-tack-toe image imposed upon your lens. Place the
subject of interest on one of the four intersections. If
you have diagonal line, it should originate on the left side
of the photograph because the eye, in our culture, reads
from left to right. If you have multiple subjects of
interest, it is best to have an odd number of subjects, as
the placement will appear less static then an even
number of subjects. Three large yellow lady’s-slippers,
Cypripedium parviflorum var. pubescens were photographed
on Dorcas Bay in the Bruce Peninsula, Ontario on
Kodachrome 25 film at 1/60 F11 with a flash. Try back
lighting or side lighting for more impact. For further tips
on composition read the excellent article on Picture Perfect
282
Orchids by the Slow Empiricist in the June 1998 issue of

the North American Native Orchid Journal at pp. 133 – 147.
More Information
Photographing wildflowers, especially native
orchids, has been an interest of mine for nearly 20 years.
An excellent way to learn more about photography is to
join a nature photography organization. I am indebted
to the Minnesota Nature Photography Club for their
many lectures and field trips through which I learned
much. Contact your local photography store to obtain
the names of your local camera clubs.
Dianne Plunkett Latham, 7013 Comanche Ct., Edina, MN 55439

E-mail: PlunkettDi@msn.com
Dianne is the former President of the Minnesota Nature Photography Club and
the past chair of the Minnesota Exhibition of Botanical Photography. She is a
three-star exhibitor in the Photography Society of America Nature Division
283
Keenan: 3RD ANNUAL NORTH AMERICAN NATIVE ORCHID
284
3RD ANNUAL NORTH AMERICAN

NATIVE ORCHID CONFERENCE
FIELD TRIPS, JULY 10 & 11, 1998
Philip E. Keenan
Lake Itasca State Park, Minnesota, is near the

dividing line between the forested and open prairie of
northwestern Minnesota, just southwest of Bemidji, a
small city of 11,000 people with a convenient air link to
Minneapolis. This allowed for a fair diversity of orchids
seen on the two days of field trips.
In a wooded bog adjacent to the park an unusual

juxtaposition of three species of adder's-mouth
orchids, Malaxis, occurred in full bloom: bog adder's-
mouth, M. paludosa, white adder's-mouth, M.
brachypoda, and green adder's-mouth, M. unifolia. The
very rare M. paludosa produced nearly one hundred stems
of perhaps the smallest terrestrial orchid flowers on the
North American continent. It was interesting to note
that here in the shade of a wooded bog the flowers were
noticeably smaller than those in an open, sunlit bog in
Anchorage, Alaska a few years ago. Large numbers of
blunt-leaved rein orchis, Platanthera obtusata, in prime
condition, were scattered among heart-leaved
twayblades, Listera cordata, most of which were past
285
prime. Many people were attracted to several good

specimens of pad-leaved orchis, Platanthera orbiculata.
Also noted were the almost ubiquitous northern

green bog orchis, Platanthera hyperborea, just another
green orchid to many, but not all, participants—who ran
the gamut from beginner to expert—and the recently
differentiated pale-green bog orchis, P. huronensis. In
the ditches, a few remaining showy lady's-slippers,
Cypripedium reginae, still in good color, provided a final
photographic opportunity, and one clump of long
bracted green orchis, Coeloglossum viride var. virescens, did
likewise. (As in many areas of the country, this spring
was two weeks earlier than usual in Minnesota). The
northern small yellow lady's-slipper, Cypripedium
parviflorum var. makasin, was one of the few orchids not
in bloom. Also seen at this site were large yellow
lady's-slipper, Cypripedium parviflorum var. pubescens,
lesser rattlesnake orchis, Goodyera repens, ram's-head
lady's-slipper, Cypripedium arietinum, small round-
leaved orchis, Amerorchis rotundifolia, and hooded
ladies'-tresses, Spiranthes romanzoffiana, which were all
present either in the woods or along the adjacent road,
For an excellent total of sixteen species and varieties.
Despite the large number of people, foot traffic was
carefully controlled, resulting in minimal damage.
The highlight in the field was the second day‘s

drive of more than three hours (each way) to extreme
southern Manitoba‘s Tall-grass Prairie Preserve near
Tolstoi, Manitoba. Why Manitoba and not the
Minnesota prairie, you ask? Because this was a poor year
286
in Minnesota where apparently it never produces the

quantity of prairie fringed orchids that Manitoba
consistently does. Perhaps 10,000 flowering, 3-foot,
sturdy stems of western prairie fringed orchid,
Platanthera praeclara, waved back and forth in moderate
breezes under a blistering sun.
The size and substance of the intensely white lips

is most impressive. For example, the middle segment of
the thrice-divided lip measures just under two inches in
diameter! This is larger than any other fringed orchid,
including the eastern prairie fringed, Platanthera
leucophaea, which it clearly surpasses in size and beauty.
The one advantage the eastern has over the western is its
sensational strong mid-day fragrance. The western offers
its fragrance, interestingly, only at dusk.
The western prairie fringed orchid is rated

second only to Cypripedium reginae as the most spectacular
North American native orchid north of Florida.
287
Philip E. Keenan, 31 Hillcrest Dr., Dover, NH . 03820

Philip is a regular contributor to the Journal and is the author of a
myriad of articles on native orchids as well as a new book, Wild
Orchids Across North America, to be published by Timber
Press this fall. E-mail: sk2325@server1.rscs.net
Platanthera praeclara
western prairie fringed orchis
288
LOOKING FORWARD
December 1998
PROCEEDINGS OF THE
RD
3 ANNUAL NORTH AMERICAN NATIVE
ORCHID CONFERENCE
Part 2
…….and more!
4th ANNUAL NORTH AMERICAN

NATIVE ORCHID CONFERENCE
April 8-13, 1999
University of South Florida

Tampa, Florida
289
Thursday & Friday - April 8 & 9
Field trips in north-central Florida
Ocala/Gainesville area
Saturday & Sunday - April 10 & 11
Conference Sessions at USF
Monday & Tuesday - April 12 & 13
Field trips in south-central Florida
Sebring/Naples area
Registration Fee of $45 per person includes all field

trips and conference sessions
A schedule of the field trips and suggested list of

accommodations will be sent upon registration
Space is limited to 75 so register early

Send your check for $45 per person to
North American Native Orchid Alliance
PO Box 772121
Ocala, Florida 34477-2121
The 1998 conference was sold out several months prior to the July dates.
Do not be one of the members who will have to be turned away for the
1999 conference. Register and make reservations early!
Last year we had several members who made their airline and hotel
reservation in December but neglected to register for the conference
until the last minute. This year when the conference is full we will not
be able to take add-ons even if you have made other paid reservations!
290
BOOK REVIEWS
BOOK REVIEWS:
The Genus Paphiopedilum, Natural History and

Cultivation
Dr. Guido Braem, Charles O. and Margaret L. Baker
1998 ISBN 0-9665337-0-4
Botanical Publishers, Inc., Kissimmee, FL
Distributed by Alpha Orchids, Miami, FL
180pp. Paper. 9x12" $49.95
47 color photographs, 3 color paintings, 71 line
drawings, 6 maps
The latest addition to the global orchid literature

is one of the most beautifully produced books of its
type. The large page format, superbly printed color
(mostly full page), illustrations and vast amount of both
botanical and horticultural information leave little room
for criticism. Although not dealing with native North
American orchids, this volume is the first of a series that
will eventually cover all species of Paphiopedilum,
Phragmipedium and related genera, and Cypripedium. The
latter will be of special interest to native orchid
enthusiasts.
The first section of this volume deals with

taxonomy—a subject that is very dear to me—and
292
BOOK REVIEWS
critical to anyone who wants to truly understand the

subject. Dr. Braem has clearly, accurately, and, at times,
in great detail, given a crash course in botanical
nomenclature and taxonomy, contrasted it to
horticultural nomenclature, and given examples of
perpetuated errors and pitfalls. He often cites both
historical and contemporary examples of both proper
and erroneous taxonomic citations. The section is an
absolute must for all native orchid enthusiasts—whether
rank amateur or professional!
The second section is a detailed history and

synopsis of the subfamily Cypripedioideae and the genus
Paphiopedilum from both the historical and contemporary
perspective. The majority of the section is devoted to a
complete listing of all species in their respective sections.
The last section is the bulk of the volume which

contains detailed accounts of 31 species (of the 89
listed), with dramatic full-page color photographs,
historical information, habitat data and just about
everything you could possibly want to know about the
culture of each individual species.
An index is provided to taxa and it will be

cumulative in each forthcoming volume. Only very few
typographical errors were found: lilly for lily and a
footnote when no footnote reference appeared. If I
could have suggested anything different it would have
been to include more in situ photographs and a Literature
Cited for each volume. Although there are copious
literature references throughout the text, the Literature
293
BOOK REVIEWS
Cited is scheduled for the concluding volume. The

remaining species of Paphiopedilum are scheduled for
Volumes 2 and 3 with a projected publication date of
late autumn 1998 for Volume 2 and spring 1999 for
Volume 3.
Although not inexpensive, it is certainly worth

every dollar for the quality of both the information and
the publication. I thoroughly recommend this to all
lovers of slipper orchids everywhere.
PMB
Wild Orchids Across North America

A Botanical Travelogue
Philip E. Keenan
294
BOOK REVIEWS
1998. Timber Press, Portland, OR

ISBN 0-88192-452-0 $39.95
320 pp., 170 color photographs, hardcover, 6x9"
Memoirs and travelogues have long been a

popular vehicle for sharing personal experiences on a
wide variety of subjects. Most natural history tomes of
this type are very general and have little specific
information to generate vivid images for the reader. Not
so with Philip Keenan's assemblage of more than two
decades of traveling throughout most of North America
in search of many of the wild orchids of that continent.
Keenan‘s descriptive prose is a fitting

complement for his brilliant full-color images of wild
orchids taken in situ from coast to coast. Having
accompanied him on many of these forays I can say that
each vignette or anecdote brings me back to the day of
discovery! For those who have not been there, his
photographs will give you a taste of what can be waiting
for you.
Both the very rare and the frequent and familiar

are treated with the same reverence and respect. For
those who have lived in the Northeast the wide-spread
pink lady's-slipper is still be to admired as much as the
rapidly decreasing fairy-slipper. Keenan takes us to such
diverse places as Newfoundland, Alaska (and its
breathtaking Kodiak Island), the pinelands of New
Jersey, and the open savannas of southeastern North
Carolina. A few days in the 'sky islands' of southeastern
Arizona yield several sought-after species. Although not
295
BOOK REVIEWS
a rabid 'life-lister' as are so many in various natural

history fields, Keenan appreciates each new species and
joins many of us in completing the study of all species
within a genus.
This volume is both an excellent addition to the

native orchidist‘s library or a perfect gift for anyone who
appreciates nature. I can think of no other book on
native orchids, or wildflowers in general, that would be
as ideal as a first volume to intrigue and infect someone
with an appreciation for the natural world around them.
PMB
296

September 1998 North American Native Orchid Journal

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September 1998 North American Native Orchid Journal

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NORTH AMERICAN NATIVE

Editor: Paul Martin Brown

The Journal welcomes articles, of any length, of both a scientific

1999 Membership in the North American Native Orchid Alliance,

This has certainly been quite a summer! With

Two recent deaths in the orchid world should be

Plans are well underway for the 4th North American

Renewals will be sent out in mid September for

Paul Martin Brown,

WHAT MAKES A GOOD ORCHIDIST?

The Slow Empiricist

At the recent 1998 NANOA Conference in

These awards got me to thinking. Orchid

photography. There are other poorly-intentioned

I have been impressed with many people I have

As I first stated, enthusiasts come in many forms

Then there are the people who are interested in

successfully grown in cultivation. They also have to have

Since my column usually champions the amateur

When I was a rank novice, I was also exposed to

come up. She is truly a light in my advance in

Another woman who showed great sense gave me

Lest you think that all amateurs who have

To go back to the professional who contributes to

comprehended before moving to a new area of

Then there are the places of learning that

Lastly, there are those who write about orchids.

To sum up, I would describe the good orchidist as

committed to the field, and who wants to see the world

PROCEEDINGS OF THE 3RD ANNUAL

PROPAGATING NORTH AMERICAN

Eight North American species of Cypripedium have

lady's-slipper, C. molle, have been grown in vitro, but

Because of their showy appearance, orchids of the

requirements can be found in Cribb (1997). In the

Orchid seeds may be germinated in vitro either

One of the main difficulties in germination of

they are dormant, and 2) remove or neutralize the

Sowing the seeds before they are dormant,

In using fully mature seeds, the seed capsule is

The oxidation of germination-inhibitors is carried

PRESENT STATUS OF PROPAGATION EFFORTS

Table 1 summarizes the current status of artificial

COMMENTS ON INDIVIDUAL SPECIES

All the species in the group I consider ―easy‖ to

grown seedlings of Cypripedium californicum to flower were

TABLE 1. Status of Propagation from Seed

Species Relatively Easy to Propagate:

Species Difficult to Propagate:

Species Presently Impossible to Propagate:

*species that have been raised to flowering from seed germinated in

[ ] = Central American species.

The group of species that I consider ―difficult‖ to

―difficult‖ group, viz., C. fasciculatum, C. passerinum, and

Among the species that I consider ―impossible‖ at

that ease of germination varies greatly from clone to

The other presently ―impossible‖ species are from

As growers gain more experience with lab-

Among North American species, Cypripedium

Allen, C., ed. 1996. North American Native Terrestrial Orchid

Cribb, P. 1997. The Genus Cypripedium. Timber Press, Inc.,

Frosch, W. 1985. Asymbiotische Vermehrung von Cypripedium

Top: Cypripedium parviflorum var. pubescens flat-petalled variant