NORTH AMERICAN NATIVE

ORCHID JOURNAL
______________________________________
Volume 4 December
Number 4 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 2
AT A LOSS?………………………………….and more!
 NORTH AMERICAN NATIVE
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.
 NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 4 December
Number 4 1998
CONTENTS
NOTES FROM THE EDITOR
297

PROCEEDINGS OF THE 3RD ANNUAL NORTH

AMERICAN NATIVE ORCHID CONFERENCE
8-11 JULY, 1998 LAKE ITASCA, MINNESOTA
Part 2
298

PLATANTHERA PRAECLARA
STRATEGIES FOR CONSERVATION AND
PROPAGATION
Margaret M. From and Paul Read
299

CYPRIPEDIUM HYBRIDS IN MAHNOMEN

COUNTY, MINNESOTA
Rob Freeman
333

AT A LOSS?
The Slow Empiricist
336

COLOR, FORM AND VARIATION

Paul Martin Brown
 343
th
4 ANNUAL NORTH AMERICAN NATIVE
ORCHID CONFERENCE
364

LOOKING FORWARD:
March 1999
368

Unless otherwise credited, all drawings in this issue are by Stan Folsom
Color Plates:
1. p. 349 Corallorhiza striata var. vreelandii; Cypripedium
reginae forma albolabium
2. p. 350 Malaxis brachypoda forma bifolia; Epidendrum
floridense
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 4, pages 297-368; issued December 31, 1998.
Copyright 1998 by the North American Native Orchid Alliance, Inc.
Cover: Goodyera pubescens by Stan Folsom
 NOTES FROM THE EDITOR

As many members are still enjoying the afterglow of the

conference this past summer, plans are well underway for the 4 th
Annual North American Native Orchid Conference in Florida this
coming April. Although the format is a bit different, and each year
probably will have a slightly different format, the conference is
nearly full. If you are planning on attending, PLEASE do not put
off registering.
This issue brings nearly all of the remaining proceedings
from the 1998 conference. Because I did not receive copy in a
timely manner, two of the major talks are not included. I do hope
that they can be a future issue. This issue is smaller than usual for
that reason, as well as fewer colored pages.
Your renewal notices are included with this issue. Please
return them as soon as possible. Several members have already
sent in their renewals without even a notice! Because my
workload is even greater now in Florida, not all of the renewals
were sent out separately. Also, I apologize for the lateness of this
issue and the lack of an index. The index will be included in
March as a separate unit.
Your continued support is appreciated as we go into our 5 th
year in 1999.

Paul Martin Brown, editor

PO Box 772121
Ocala, Florida 34477-2121
naorchid@aol.com
Telephone & fax: 352/861-2565

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 PROCEEDINGS OF THE 3RD ANNUAL
NORTH AMERICAN NATIVE ORCHID
CONFERENCE
8-11 JULY, 1998 LAKE ITASCA,
MINNESOTA
Part 2

Platanthera praec1ara, a threatened prairie orchid

Margaret From & Paul Read
Color, Form and Variation
Paul Martin Brown
Cypripedium hybrids of Mahnomen County, Minnesota Rob
Freeman

To appear at a later date:

Recent Advances in the Systematics and Ecology of North
American Orchids
Dr. Paul M. Catling
Recent Research on Minnesota Orchids
Welby Smith

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AND PROPAGATION

Strategies for Conservation and Propagation

Margaret M. From and Paul Read

Introduction

The Western Prairie Fringed Orchid, Platanthera

praeclara, is listed as a Threatened species, and as such, is
afforded protection under federal and state Endangered
Species acts. Federal and state permits were obtained to
conduct this research. The species has resisted attempts
at propagation when using traditional propagation
methods, according to the Nebraska Game and Parks
Commission (Fritz, 1993). Integrated conservation
strategies combining micropropagation techniques,
histological studies, and in-situ species management are
being employed for P. praeclara, whose population
numbers are believed to be in decline. Several issues are
being investigated in this ongoing study: 1) protocols for
germinating P. praeclara seeds in-vitro, 2) the study of seed

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structures and their possible effects on germination
responses in-vitro, 3) histological studies of in-vitro
produced P. praeclara tissues using scanning electron
microscopy and conventional high magnification
techniques and, 4) a limited hand pollination study to
assess the possibility of inducing greater fruit set within a
wild population.

Seed Germination Response

Mature seeds collected in late summer of years 1995 and

1996 were air dried for five days in the lab, removed
from the fruits and placed in sealed glass vials under
refrigeration until they were cultured. Seeds were
surface sterilized in one of two solutions; an 8% calcium
hypochlorite or a 10% sodium hypochlorite solution,
and rinsed in sterile distilled water before being
aseptically cultured on agar-gelled media.

The water repellent seed testa is dense and bleaching

removes much of the brown pigmentation (Stoutamire
1981). Frequently this occurs earliest at the suspensor
end of the seed during the surface sterilization process.
This could be a factor in the damage caused to that polar
region when the sterilization solution is too harsh or
applied for too long. The carapace surrounding the
embryo cells also appears dark and dense, with no visible

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opening. This creates a secondary hydrophobic barrier,
and can further restrict embryo water uptake.

Germination responses were obtained on a

modified Fast medium (Fast 1982), with additional
modifications (Anderson 1990), a 1/3 strength MS
(Murashige and Skoog 1962) as recommended (Chu and
Mudge 1994), amended with 40ml. L coconut water and
6 g.L agar, and a new medium designated as P/C (From,
unpublished).

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Cultures were incubated for 6 weeks at room
temperature, 23 C ± 2, in darkness. Cold treatments
were applied for 30, 60 or 90 days respectively for each
of the 3 annual seed sowings. Thirty days of cold
stratification resulted in a lower germination response
than either a 60 or 90 day cold treatment. After the cold
treatments, cultures were returned to laboratory room
temperatures, still in continual darkness, until
protocorms developed a root initial and a shoot initial.
Protocorms which developed shoots 2 mm in height or
greater, were placed under cool white fluorescent lights
for the remainder of the natural growing season.

Initial germination began after 150 days with 1995

seeds and after as little as 18 days with 1996 seeds.
However, responses were highly variable between
individual cultures and only a few cultures displayed 18
day germination response. A small number germinated
early, but in-vitro sown P. praeclara seeds do not easily
synchronize, and germination continued for up to 17
months within some vessels.

Seed Structures

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Mature seeds were photographed both in the dry state,
and after soaking in a bleach solution, using a Nikon
FX-35 DX camera mounted on a Nikon Labophot-2.
Seeds were also sputter coated with gold and
photographed using scanning electron microscopy
(SEM), on a Philips 515. Elemental analysis was
performed with a Kevex 7000.

Dry mature seeds display a dark, opaque testa,

with a distinctive, reticulate pattern unique to the
species. The testa provides a primary barrier to water
uptake necessary for the embryo's cell division during
germination. The enclosed bare embryo is surrounded
by a well-developed, dark carapace which is
hypothesized to provide a secondary barrier to water
uptake, (figure 1). No visible opening is apparent in the
carapace, unlike seeds of the European native orchid,
Orchis morio, whose seeds are reportedly easy to
germinate in-vitro (Ronse, 1989). Seeds soaked in bleach
solution were photographed over a 2 ½ hour period to
record changes in the testa or embryos. Soaked seeds
displayed a slightly more transparent seedcoat, which in
some instances showed liquid collecting at a kevelian
border momentarily, and then rapidly sheathing the
space between and collecting at another border, (figure
2). As the duration of soaking time increased, the liquid
appeared to flow past the embryo with progressively
greater ease. Repeated experimentation is necessary to

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determine the optimal soaking time to adequately soften
the seedcoat, without doing serious damage to the
immature embryo, in order to maximize germination
response. Physical dormancy may be influenced by a
number of factors, including natural environmental
stresses during fruit maturation, genotypic variance, soil
types, plant nutrition, length of growing season,
precipitation, dehiscence, seed storage techniques, and
other, still unidentified dormancy factors.

Scanning electron microscopy was used to record

the seedcoat pattern, (Appendix I, fig. 1.1). A semi-thin
section of a single seed displays a total of 30 individual
embryo cells. The section was stained with Hematoxylin
and Eosin, (H & E), which stains living cells pink and
leaves the dead, brown cells of the broken testa
unstained. No external endosperm is visible, (Appendix
I, fig. 1.3a). Individual embryo cells do contain small
spherical bodies. When those bodies are magnified to
836X SEM, (Appendix I, fig. 1.3b), elemental analysis
shows that they consist primarily of potassium.

Platanthera praeclara
Plant Tissue Histological Study

Little information is available about the basic

biology or culture of P. praeclara. One concern

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frequently voiced by biologists and conservationists is
that any species propagated for reintroduction efforts
must be true to the wild genotype. Consequently, in-vitro
produced plant tissues are being photographed for plant
developmental studies. When the same P. praeclara seed
semi-thin section, noted above, is photographed on a
conventional microscope, a small arc of yellow-gold cells
are visible at the suspensor end of the embryo. It is
hypothesized that this region is the area of most rapid
cell division during germination. (Appendix I, figure 1.4)

Figure 1.5, Appendix I, is a semi-thin section

showing considerable cell division by the time the
protocorm reaches 2 mm in length. Meristermatic
regions contain the densest concentration of cells.
Individual protocorm cells at 1930X SEM reveal clusters
of bodies which are hypothesized to consist of mobilized
proteins and starches, (Rasmussen, 1995). These
fracture when scanned, leading one to conclude that
these bodies consist of soft tissue. Staining the semi-
thin protocorm section results in black-stained nuclei.
The young seedling appears to have coalescing proteins
which contribute to the formation of starch grains,
producing an increased total cell volume, (Appendix I,
fig. 1.6)

After 10 months, some P. praeclara protocorms

from 1995 began to develop a shoot initial, a root initial

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and a structure resembling a small tuber. However,
tuber-like structures in the first year of aseptic culture,
appear to be the exception rather than the rule. The
majority of protocorms develop hair-like rhizoids
protruding from the surface of the spherical protocorm.
Protocorms which display exceptional vigor can develop
2 to 4 roots up to 12cm in length by the end of 15
months. Tubers develop into a variety of shapes in-vitro,
which closely resemble those found on plants excavated
at the Sheyenne National Grasslands, (Wolken 1995).
Tubers may be coiled, rod-shaped, oval or tapered,
(Appendix I fig. 1.7). Alternating cold treatments with
room temperature regimes annually appears to have a
favorable impact on tuber formation, particularly with
those protocorms which had at least one shoot 5-20mm
in height. Protocorms not given an annual cold
treatment frequently become necrotic after 12
continuous months at room temperature.

Hand Pollination Study

Two orchid sites were chosen to conduct human-

assisted pollination. Site #1 is an upland site on a
privately-owned prairie and site #2 is a wet, low-lying
prairie swale on federal lands. The sites are located
approximately 350 miles apart.

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The study was first conducted in 1996. Its
primary purpose is to assess whether there is any merit
in using human assisted pollination to improve seed set
for the orchid genotype(s) in wild populations located in
the far western reaches of the orchid's range, where it's
populations are small, under threat of encroachment and
the orchid population numbers are believed to be in
decline.
Twenty-eight plants were randomly chosen for study at
site #1 and 10 plants were randomly chosen at site #2.
Each individual inflorescence displayed a minimum of 5
fully- expanded, intact flowers. At the upland site; site
#1, eight plants were cross pollinated with another
individual plant a minimum of 20 meters away. Four
plants were self-pollinated, and seven plants receiving no
human assisted pollination were treated as a control
group. All plants in the study were tagged
inconspicuously, staked and recorded. At the prairie
swale site; site #2, located on federal lands, permission
was obtained to hand pollinate five plants and five
additional plants were marked as control plants.

Pollen sacs were removed from one individual

flower with the aid of toothpicks and placed on the
stigma region of another flower. Manipulations and
staking were completed at site #1 on June 28th, 1996.
Anthesis commenced 19 days later at site #2, and hand
pollination at that location took place on July 17, 1996.

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Data were gathered at site #1 on September 13,

1996, and results indicated a much higher average fruit
set on plants which were cross-pollinated, 5.1 per plant,
than on selfed plants, 2 per plant, or control plants, 1 per
plant, at site #1. Fruit set on cross pollinated plants 5.3
per plant, versus fruit set on control plants 2.7 per plant,
followed a similar pattern at site #2 in 1996. Due to the
extensive logistical and legal limitations placed on the
experiment at each of the two sites in 1996, the greatly
reduced numbers of flowering individuals available at
site #1 in 1997, and the adverse weather conditions at
site #2 in the 1997 growing season, data are presented
here as a starting point for research in the area of human
assisted pollination to increase fruit production. More
research is needed before any recommendations should
be made whether this practice is beneficial for long term
species conservation. Preliminary data indicate that
human assisted pollination may indeed increase fruit set
in a wild population. Caution must be exercised since
seed production is a costly function in terms of most
species' total energy reserves. Plants which were cross-
pollinated also remained in a vegetative state much later
into the autumn, and it is currently unclear whether
delayed dormancy may affect an individual plant's
longevity. Moe and Pleasants in 1993 also questioned
whether the recorded fruit set levels they studied were

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adequate to maintain the species at a given site. Further

research is needed to adequately answer these concerns.

Assessment is needed of the threat of extinction to

current populations of the Western Prairie Fringed
Orchid. Future research into its propagation for
possible reintroduction efforts should recognize several
important factors: 1) the possibility that a single threat
could be capable of causing 100% mortality in a
sufficient number of the very small populations (those
with 10 or fewer plants) scattered throughout the state,
so as to pose a real threat to the species' viability within
that state or region, 2) the likelihood of natural
fluctuations in populations to synchronize at a low point
that could threaten the species' viability, 3) the genetic
viability and variability of the species in a given region,
and 4) the likelihood that suitable habitat will continue to
be available. These issues are particularly important in
areas where intensive cultivation practices and their
accompanying heavy uses of herbicides, pesticides and
fertilizers, or heavy grazing practices and repeated
mowings are carried out, each of which can dramatically
effect WPFO fruit-set.

Although simultaneous floods, fires or disease

pose a relatively small threat to all populations
throughout the orchid's range, one population may

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represent a considerable percentage of the species'
genetic base in sparsely populated regions. Therefore,
the loss of even one, or a few small populations
simultaneously, could drastically reduce the species' total
genetic diversity. It is currently unknown how much
genetic exchange occurs by natural pollen vectors among
the individual sites, and future research may point to the
possibility that manipulated crossings between
populations may be necessary to maintain species
viability, since populations are now often found in non-
contiguous colonies. One possible future remedy may
be to reintroduce new populations in protected areas
within the species' historic range.

The occurrence and magnitude of the species may

overall be relatively high, particularly in the northeastern
portions of its range. However, this may be of little
consequence in the western and southern portions of its
range if individual populations do not remain viable over
the long term, and diminished populations no longer
have any genetic exchange between their disjunct
locations. Propagation as a possible source of WPFO
plants in the future, may become increasingly important
for the species' preservation and continued presence in
the western portions of the orchid's range.

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Acknowledgments

Partial funding was provided by:

Nebraska Environmental Trust
Iowa Living Heritage Roadway Trust
Nebraska Statewide Arboretum
Mid America Orchid Conference
Association of Zoological Horticulture
Omaha Henry Doorly Zoo

Gratitude is extended to the following for their

technical or logistical assistance: UNL Institute of
Agricultural and Natural Resources, Marty Cano at the
University of Nebraska Medical Center in Omaha for
SEM technical assistance, and Len McDaniel for field
assistance from the U.S. Fish and Wildlife Service. Dr.
Lee Simmons, Director and Terri Gouveia, Horticulture
Curator at Omaha's Henry Doorly Zoo provided
invaluable assistance. Virginia Miller, UNL technologist,
and the Nebraska Game and Parks Commission
contributed technical and field insights. Karen Delaney
provided histological assistance at UNMC. Drs. Karen
Johnson of University of Manitoba's Museum of Man
and Nature, Devonian Botanic Garden, University of
Alberta, and Marlin Bowles at the Morton Arboretum
provided mycelium cultures for symbiotic study. Drs.
Charles Sheviak, Warren Stoutamire and William Steele,
as well as Margaret Ramsey at Royal Botanical Garden

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Kew, each were kind enough to provide advice and
encouragement.

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Literature Cited

Anderson, A.B. 1990. Asymbiotic germination of seeds

of some North American orchids, in C. E. Sayers
{ed.} North American native terrestrial orchid
propagation and production. Brandywine
Conservancy, Chadds Ford, Ps. Pp. 75-80

Chu, C. C., and K. Mudge. 1994. Effects of prechilling

and liquid suspension culture of seed germination
of the yellow lady's slipper orchid (Cypripedium
calceolus var. pubescens). Lindleyana 9(3): 153-159

Fast, G. 1982. European terrestrial orchids - symbiotic

and asymbiotic methods. In: Orchid Biology,
Reviews and Perspectives II. J. Arditti [ed.].
Comstock Pub. Associates. Ithaca, N.Y. pp. 309-
326.

Federal Register. 1989. Rules and regulations. 54: 160-

167.

Murashige, Toshio. 1974. Plant propagation through

tissue cultures. Ann. Rev. Plant Physiol. 25: 135-
66

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Pleasants, J. And S. Moe. 1993. Floral dispay size and

pollination of the western prairie fringed orchid,
Platanthera praeclara (Orchidaceae). Lindleyana 8
(1): 32-38.

Rasmussen, H. 1995. Terrestrial Orchids from Seed to

Mycotrophic Plant. Cambridge Press, U. K. pp.
28-34

Ronse, A. 1989. In vitro propagation of orchids and

nature conservation: possibilities and limitations.
Mem. Soc. Roy. Belg. 11: 107-113

Stoutamire, W. 1981. Early growth in North American

terrestrial orchid seedlings. Pages 14-24 in E.H.
Plaxton [ed.] Proc. Symp. II and Lectures.
Southfield, Michigan.

Wolken, Paige M. 1995. Habitat and life history of the

western prairie fringed orchid (Platanthera
praeclara). Thesis to University of Wyoming.

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Fig. 1.1 An individual seed of Platanthera praeclara

photographed by scanning electron microscopy (SEM).
The testa displays the characteristic reticulate pattern
unique to this species. Seeds of terrestrial orchids each
have their own distinct pattern. The microscopic seed
was photographed at 160x.

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Fig. 1.2 A longitudinal semi-thin section of P.

praeclara seed stained with Hematoxylin and Eosin,
(H+E). Living embryo cells stain pink. The dead brown
cells of the testa remnants do not stain. Photographed
at 140x on a Nikon Labophot FX35.

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Fig. 1.3 (a) P. praeclara seed photographed in a

longitudinal semi-thin section using SEM at 312x. The
individual cell walls within the embryo are visible (1) and
fragments of the dissected testa remains (2). (b) Bodies
within the seed section from (a)enlarged to 836x. (c)
The photograph on the lower right illustrates the
location of those bodies in (b), within one seed cell, at
625x. The highlighted rectangle is enlarged from (b).
Elemental analysis of these bodies, using a Kevex 7000,
identifies them as consisting primarily of potassium.
Nutrient reserves are visible within the seed embryo
itself but none are visible in the area between the
embryo and the testa.

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Fig. 1.4 The same P. Praeclara seed as in Fig 1.3

photographed on a Nikon Labophot FX-35 at 250x.
The yellow-gold cells (1) at the suspensor end of the
embryo represent the most actively dividing cells during
germination. The testa's dead cells have been broken (2)
by the sectioning process.

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Fig. 1.5 (a) A semi-thin section of a P. Praeclara

protocrom by SEM at 30X. Cell density is greatest at the
apical meristem end (1). Nuclear volume has increased
as the cells enlarge and divide. Cell density at (2) may by
the earliest signs of another stele or a root meristem in
an early developmental stage. (b) The same protocorm
interior at 312x SEM. Individual nutrient bodies (3)
within cells are visible. © Those same nutrient bodies
within the highlighted rectangle from Fig. 1.7, (b) at
1930x SEM. Elemental analysis of those bodies
indicates that they consist primarily of potassium and
calcium. Calcium is a component of cell walls, and the
analysis may be picking up the surrounding walls.

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Fig. 1.6 The protocorm semi-thin section

photographed at 200x after staining with H&E.
Hematoxylin stained the nuclei black (1) and all other
living tissue is stained pink. Coalescing protein
vacuoles (2) and newly formed starch grains (3) show
that the germinated seedling is capable of mobilizing
stored protein bodies to increase cell volume as they
enlarge and divide.

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Fig. 1.7 Protocoms removed after 19 months in

asymbiotic cultures. Tubers resemble those excavated
from the natural environment by researchers at the
Sheyenne National Grasslands (Wolken 1995).

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Fig. 1.8 P. praeclara root surface. The surfaces

show fungal hyphae infection after 6 weeks in
symbiotic culture. SEM at 573x.

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Fig. 1.9 A cross-section of root cells which had not

made contact with mycelium inoculated onto the
substrate. Individual cell walls are visible. SEM 680x.

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Fig. 1.10 Infected root cells containing bodies

resembling starch grains. SEM at 1490x. When scanned
individually, they indicate soft tissue.

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Fig. 1.11 Longitudinal section taken of root infected

cells in symbiotic culture. SEM at 225x.

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Fig. 1.12 Cross section of the orchid root with

fungal hyphae inter-connecting individual root cells.
573x SEM.

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Fig. 1.13 Infected orchid root and rhizoids in cross-

section at 252x SEM. Rhizoids are single-celled
extensions of epidermal hairs. They appear as hollow
tubes.

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Fig. 1.14 Cross-section of infected root at 60x SEM.

Note the congested appearance of root cells which had
made contact with the mycelium in the symbiotic
culture.

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Fig. 1.15 Tropical orchid seed which exhibits openings in

the testa. The seeds of this species germinate easily in
asymbiotic culture. 1640X SEM

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Margaret M. From and Paul Read

Horticulture Department
University of Nebraska Lincoln
Lincoln, NE USA
Omaha Henry Doorly Zoo
Omaha, NE USA

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CYPRIPEDIUM HYBRIDS IN
MAHNOMEN COUNTY, MINNESOTA

Rob Freeman

Mahnomen County, located in North-West Minnesota,

may be home to the largest population of hybrids between
both small white lady's-slipper, Cypripedium candidum and northern
small yellow lady’s slipper, Cypripedium parviflorum var. makasin. Also
there is some indication that the large yellow lady's-slipper,
Cypripedium parviflorum var. pubescens may have hybrid with Cypripedium
candidum, but not as evident as the aforementioned. The majority
of the orchids were located in a MN-WMA (Wildlife
Management Area); which is a protected area from the plow
or other construction.

The sheer number of orchids present along with the

many variances among the hybrids made it very difficult to
distinguish exactly which species crossed with which species.
A rough estimate as to how many plants of anyone given
species were present is about one plant per every 16 square
feet on average. Some areas were so dense that it was literally
impossible not to step on a plant. The estimates given are for

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flowering plants only, I'm quite positive there are just as many
non-flowering plants if not more.

The area is predominantly open prairie with aspen

(Populus tremuloides) wooded islands forming around prairie
potholes. The transition zone between prairie and woodland
consists primarily of Silver Buffaloberry (Shepherdia argentea). It is
within this transition zone that the majority of exotic hybrids
occur. I have seen plants with a yellow lip that was almost
completely blotched out in red, much like that of spotted
lady's-slipper, Cypripedium guttatum. On several occurrences the
lips were grotesquely deformed, possibly by a late frost.
The whole floral structure on most of the species within
this zone were also not much bigger than a dime.
Another major difference that I noticed was the behavior
of the dorsal sepal. The sepal would bend back, away
from the lip at a 90 degree angle, the interesting point
about these plants is the fact that this occurrence is
random among the population and each individual
flower within the plant had the exact same properties.
The peak flowering time was June 13th in 1997, and will
vary with the weather. When I was there on the 13th of
June I noticed a short, large flowered hybrid just
beginning to bloom on the fringe of the open prairie and
brush. These are just several examples of the more
prominent hybrid features I noticed, there were many
more to numerous to mention.

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Freeman: CYPRlPEDIUM HYBRIDS IN MAHNOMEN CO., MN

The county with its vast array of Cypripedium's is an

excellent area to explore and study. The roadsides are filled
with Cypripedium candidum and do not seem to be phased by minor
disturbance. The plants of C candidum seem to benefit by slight
disturbance, for instance in an area where a bull-dozer did
some work the orchid seems to be more prolific than the
surrounding areas.

Rob Freeman, 98 8th St N, Sartell, Minnesota 56377

FreemanR@mn-amg.ngb.army.mil

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 Brown: COLOR, FORM, AND VARIATION

AT A LOSS?

The Slow Empiricist

When I started to read the articles in preparation

to format them for publication in the current issue of the
North American Native Orchid Journal marveled at the
scientific discipline and dedication the authors
exhibited. These people use the latest techniques to
investigate their subjects, employing microscopy that
magnifies things to amazing degrees as in Margaret
From's article. They employ careful scientific
measurements to document their findings and adhere to
rigid principles. As a proponent of the novice orchid
enthusiast I wondered what I could possibly write that
would even come close to such learned works. I still
believe that the novice and amateur bring a certain zeal
and refreshingly different point of view to the orchid
world that is worth fostering. The winter months in the
Northern Hemisphere bring a halt to much of the
outdoor activity that many amateurs enjoy in their
orchid pursuits unless they live in the extreme southern
limits of the hemisphere. This is no time, however, to
cease in your activities. As I have urged in previous
articles you can find information in books and through

336
 Brown: COLOR, FORM, AND VARIATION

course-work to increase your stores of knowledge. This

article is about what you may find in nature if winter has
ended the growing season in your part of the world.

The northern parts of the North American

continent where I lived for many years have entered the
winter phase of their year with frozen landscapes and
chills in the air. Growing things have ceased to put on
new growth and there is a dearth of flowering material to
be seen out in the wild. This puts those who like to trek
about in nature finding flowering plants at a distinct
disadvantage. Many such enthusiasts, especially the
novices, think there is little out there to entice them to
leave their cozy, warm homes. The orchid hounds, who
only want to look at orchids, might feel at a real loss
about how to f1ll their need for orchid expeditions to
view their favorite quarry under these circumstances.

Contrariwise, there are many who know of the

joys of experiencing nature even in the dead of winter. If
you are not so bound by your love for orchids in bloom
that nothing else will make you happy, you can take a
walk in the wilds and find many things of interest, even
some orchids. However, they won't be in flower in the
northern climes. You will have to travel to the southern
parts to enjoy blooming specimens.

In November, for instance, in the northern New

England area, one can set out on a sunny, dry morning
and wander into mesic forests and find evidence of things

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 Brown: COLOR, FORM, AND VARIATION

still growing and some things in bloom. Witch up more

lasting evidence of their existence. Some signal their
presence with a hardy seed stalk that will last through the
winter and even into the next blooming period. Others
have winter rosettes that attract the eye.

You should be able to find seedpods of many

orchids that were successful in attracting a pollinator and
setting seed. These hardier plants than the tender or
fragile varieties often but not always keep their seed
stalks into the next year and display them beside their
new blooming stems. I have usually found plants of the
pink lady's-slipper, 0pripedium acaule, in the Maine woods
in winter or early spring by their prominent seed stalk
standing proudly above the forest floor. It has a medium
brown colored stalk and seed pod. It stands straight up
about 12 inches (mm) with a slightly inclined seed head
that has several compartments sheathed in a sepal-like
enclosure. The large whorled pogonia, Isotria verticillata,
and its cousin, the small whorled pogonia, Isotria
medeoloides both put up a sturdy seed pod much like the
Cypripediums except that its seed pod stands straight up on
the top of the stem. It is also shorter than the Cypripedium
stalks. These all can be encountered if you are open to
the possibility and can be found if you are in the right
territory. The rattlesnake. orchids, Goodyera pubescens, G.
tesselata, G. repens, and G. oblongifolia have spikes that
parade a series of swollen seedpods along their stems.
Even the more fragile coralroots can withstand the
ravages of nature to leave a marker that they existed. If
you know what the fruiting stage of orchids looks like
you should be

338
 Brown: COLOR, FORM, AND VARIATION

Hazel will have clusters of bright yellow flowers

clinging to the branches because it is a November
bloomer. They are not very large and you might miss
their display but if you find them they will brighten
your expedition. Other species of witch hazel will
bloom in gardens later in February, or, the early spring
with orange-yellow blossoms. These put on a more
spectacular show.

Holly is another plant that puts on a cheery

display with its bright red berries and glossy leaves.
The deciduous varieties tend to grow in swampy areas
where you can get your shoes damp to downright wet.
If you bring along a pair of clippers you might be able
to prune a few stems for your empty window boxes.
They look splendid with a few evergreen boughs for a
background and will last for some time. There may also
be some of the red-orange berries of the bittersweet
vine to treat your eyes as you ramble in your quest to
find evidence that there is life left in the seemingly
barren vistas. These berries are not very hardy and drop
quickly from their branches, so they don't make such a
lasting display as holly can.

But what of orchids I can hear you questioning.

As I stated there won't be any in bloom but they have
not all coalesced into nothingness. Some have, of
course, like the shy three birds orchid, Triphora
trianthophora. They are scarcely out when they disappear
from sight back under the leaves that litter their
habitats. At least they do in the northern areas of the
United States like New Hampshire. Other orchids put

339
 Brown: COLOR, FORM, AND VARIATION

able to spot these specimens in your expeditions. If you

don't have enough knowledge of this stage in the orchids'
growth cycle you should study the literature to find
photographs or drawings that will show you this phase
of the orchid. Blanch Ames' drawings show this stage
quite clearly.

Another way to find an orchid population is from

their winter rosette. The aforementioned Goodyeras have
beautiful rosettes with markings that make them easy to
identify. I will grant you that the smaller varieties like
Goodyera repens are much harder to spot but if you have a
keen eye and have it in the back of your mind when you
are exploring you may just be surprised to find some.

Lastly, I will mention you might find three very

rare orchids in the northern climes by their distinctive
winter leaves. They do not form rosettes but tend to put
up single leaves as evidence of their being present. The
three orchids are Calypso bulbosa, the fairy slipper orchid,
Tipularia discolor, the crane fly orchid and the puttyroot
orchid, Aplectrum hyemale. Each puts up a very distinctive
basal leaf that stands out among the brown and golds of
the dried plants that have succumbed to the winter frosts.
The leaf of Calypso can be seen rising from the sphagnum
and cedar needle littered duff that shares its habitat in
northern cedar swamps. It has a wrinkled surface and a
curving stem that holds it above the level of the duff like
a tiny flag. The leaf edges are slightly fluted or
scalloped. These

340
 Brown: COLOR, FORM, AND VARIATION

specimens can be very tiny so you need a sharp eye to

locate them. The leaves of the Tipularia are about five
inches long (12.5 cm) triangular shaped and has a
spotted upper surface that is a rich green with magenta
spots. The under surface is a brilliant magenta purple.
These leaves seem to lie on the forest floor like they
had fallen from some taller area unlike the Calypso
leaves that rise perkily from the ground on their curved
stems. If you locate these flatter leaves, mark the
location very well because the leaves will all dissipate
before a blooming stalk appears which could render
refinding them in the summer a virtually impossible
task. Aplectrum leaves are ovate and larger than the
Tipularia but they have a dull gray green upper surface.
Their undersides are a dark magenta much like the
Tipularia leaves. Much like Tipularia these leaves lie flat
on the ground and also dissipate before a flowering
stalk appears so keep a good record of where these
plants occur as well.

If you inhabit more moderate climates you may

find these orchids in more abundance. Tipularia grows
on Long Island in New York State and as far south as
Florida with larger stand in places like the Great
Smokies of North Carolina. The Eastern fairy slipper,
Calypso bulbosa, prefers the far reaches of northern
Vermont and similar states into Canada and north to
Alaska. The Western fairy slipper can be found in the
redwood forests of California. The Eastern fairy slipper
is hard to find because of its tiny size and scarcity. The
Western fairy slipper seems to be more abundant and
should therefore be easier to spot. The

341
 Brown: COLOR, FORM, AND VARIATION

other two, Tipularia and Aplectrum are easier to spot in

leaf but they still become illusive because of the
disappearance of the basal leaf before they bloom.
I wish you good luck with your explorations and
I realize you have to have a reasonably open winter to
locate these terrestrials. You can still enjoy the outdoors
even with lots of snow if you would be willing to look
at other aspects of nature, which I alluded to, in the
opening paragraphs. Let the orchids rest under their
blanket of snow.

Your Slow Empiricist

342
 Brown: COLOR, FORM, AND VARIATION

COLOR, FORM AND VARIATION

Paul Martin Brown

When does a pink lady's-slipper not look like a pink lady's-

slipper? When it has white flowers or two pouches! Many of
our native orchids can be highly variable in both color and
form. Over the years many of these variations have been
recognized at several taxonomic levels - species, subspecies,
varieties and at the forma level. I wish to review many of these
variations and discuss the appropriate level of recognition.

This paper does not intent to cover generic transfers, nor

does it intended to include all of the taxa that fall under the
respective categories. References are given for most recent
treatments of the taxa.

Those species that have been recognized at both the species

and varietal levels and are deemed to be valid species.

Some examples of this level of recognition are:

Platanthera grandiflora (Bigelow) Lindley

LARGE PURPLE FRINGED ORCHIS
Synonym: Platanthera psycodes (Linnaeus) Lindley var.
grandiflora (Bigelow) A. Gray

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 Brown: COLOR, FORM, AND VARIATION

Stoutamire, W.P. 1974. Brittonia 26: 42-58.

Cleistes bifaria (Fernald) Catling & Gregg

UPLAND SPREADING PO GONIA
Synonym: Cleistes divaricata (Linnaeus) Ames var. bifaria
Fernald
Catling, P.M. & K.B. Gregg. 1992. Lindleyana 7(2): 5773.

Cypripedium yatabeanum Makino

YELLOW SPOTTED LADY'S-SLIPPER
Synonym: Cypripedium guttatum var. yatabeanum (Makino)
Hultén
Brown, P.M. 1995. NA Native Orchid Journal 1 (3): 199.

Platanthera brevifolia (Greene) Kranzlein

SHORT-LEAVED REIN ORCHIS
Synonym: Platanthera sparsiflora (S. Watson) Schlecter var.
brevifolia (Greene) Luer

Platanthera huronensis (Nuttall) Lindley

PALE GREEN BOG ORCHIS
Synonym: Platanthera hyperborea (Linnaeus) Lindley var.
huronensis (Nuttall) Luer

344
 Brown: COLOR, FORM, AND VARIATION

Platanthera macrophylla (Goldie) P.M. Brown

GOLDIE'S PAD-LEAVED ORCHIS
Synonym: Platanthera orbiculata (Pursh) Lindley var.
macrophylla/a (Goldie) Luer
Reddoch, A.H. & J. M. Reddoch 1993. Lindleyana. 8(4): 171-
188.

P Platanthera purpurascens (Rydberg) Sheviak & Jennings SHORT-

SPURRED BOG ORCHIS
Synonym: Platanthera hyperborea (Linnaeus) Lindley var.
purpurascens (Rydberg) Luer
Sheviak & Jennings. 1997" NA Native Orchid Journal 3(4): 444-
449.

Ponthieva brittoniae Ames

MRS. BRITTON'S SHADOW-WITCH
Synonym: Ponthieva racemosa (Walter) C. Mohr var. brittonae
(Ames) Luer
McCartney, c.L., Jr. 1995. NA Native Orchid Journal 1(2): 106-
116.

Pseudorchis straminea (Fernald) Soó

NEWFOUNDLAND ORCHIS
Synonym: Pseudorchis albida (Linnaeus) Love & Love subsp.
straminea (Fernald) Love & Love
Reinhammar, L. 1995. Nordic Journal of Botany 15(5): 469481.
- 1997. NA Native Orchid Journal 3(4): 407-425.

345
 Brown: COLOR, FORM, AND VARIATION

Spiranthes. floridana (Wherry) Cory

FLORIDA LADIES'- TRESSES
Synonym: Spiranthes brevilabris Lindley var. floridana

Spiranthes ochroleuca (Rydberg) Rydberg

YELLOW LADIES'-TRESSES
Synonym: Spiranthes cernua var. ochroleuca
Sheviak, C.J. 1991. Lindleyana 6(4): 228-234.

Spiranthes odorata (Nuttall) Lindley

FRAGRANT LADIES'-TRESSES
Synonym: Spiranthes cernua var. odorata (Nuttall) Correll
Sheviak, C.J. 1991. Lindleyana 6(4): 228-234.

Malaxis brachypoda (Gray) Fernald

WHITE ADDER'S-MOUTH
Synonym: Malaxis monophyllos (Linnaeus) Swartz. var.
brachypoda (A. Gray) Morris & Eames

Those taxa that were described as species, never received

synonymy as varieties and were merged by authors into
another species.

Some examples would be:

Cypripedium kentuckiense C.F. Reed

KENTUCKY LADY'S-SLIPPER
Atwood, J. T. Jr. 1984. AOS Bulletin 53(8): 835-841. Brown,
P.M. 1995. NA Native Orchid Journal 1 (3): 255. Reed, C.
1981. Phytologia 48(5): 426-428.

346
 Brown: COLOR, FORM, AND VARIATION

Malaxis bayardii Fernald

BAYARD'S ADDER'S-MOUTH Catling,
P.M. 1991. Lindleyana 6(1): 3-23.

The last grouping at the species level would be those

species recently described that have been segregated from
existing species. These have not been reduced to synonymy
by other authors.

Calopogon oklahomensis D .H. Goldman

OKLAHOMA GRASS-PINK
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 133.
Goldman, D.H. 1995. Lindleyana 10(1): 37-42.

Epidendrum floridense Hágsater

FLORIDA UMBELLED EPIDENDRUM SYN:
Epidendrum difforme Jacquin in part
Neolehmannia difformis (Jacquin) Pabst
Hágsater, E. & G. Salazar. 1993. Icones Orchidacearum
Romero, G.A.1994. A.O.S. Bulletin 63(10): 1168-1170.

Malaxis wendtii Salazar WENDT'S ADDER'S-MOUTH

Salazar, G. 1993. Orquidea (Mex.) 13(1-2): 281-284.

Piperia candida Morgan & Ackerman

SLENDER WHITE PIPERIA
Morgan, R. & J. Ackerman. 1990. Lindleyana 5(4): 205211.

347
 Brown: COLOR, FORM, AND VARIATION

Piperia colemanii Morgan & Glicenstein

COLEMAN'S PIPERIA
Morgan, R. & L. Glicenstein. 1993. Lindleyana 8(2): 89-

Piperia yadonii R. Morgan &]. Ackerman

YADON'S PIPERIA
Morgan, R &]. Ackerman. 1990. Lindleyana 5(4): 205211.

Platanthera pallida P.M. Brown

PALE FRINGED ORCHIS
Brown, P.M. 1993. Novon. 2(4): 308-311.

Platanthera praeclara Sheviak & Bowles

WESTERN PRAIRIE FRINGED ORCHIS
Sheviak, C.J.& M. Bowles. 1986. Rhodora. 88: 267-290.

Platanthera zothecina (Higgins & Welsh) Kartesz & Gandhi

CLOISTERED BOG ORCHID
Higgins, L.C. & S. L. Welsh. 1986. Great Basin
Naturalist. 46: 259.

Spiranthes casei Catling & Cruise var. casei

CASE'S LADIES'- TRESSES
Catling, P.M. & ].E. Cruise. 1974. Rhodora 76: 256-536.

348
 Brown: COLOR, FORM, AND VARIATION

Above:
Corallorhiza
striata var.
vreelandii
Marin Co., CA

Right:
Cypripedium
reginae forma
albolabium
Orange Co., VT

P.M. Brown

349
 Brown: COLOR, FORM, AND VARIATION

Above:
Malaxis brachypoda
forma bifolia
Windsor Co., VT

Right:
Epidendrum
floridense
Collier Co., FL

P.M. Brown

350
 Brown: COLOR, FORM, AND VARIATION

Spiranthes delitescens Sheviak

CIENEGAS LADIES'-TRESSES
McClaren, M.C 1996. NA Native Orchid Journal 2(2): 151-169.
McClaren, M.C. & P.C Sundt. 1992. Southwestern Naturalist 37:
299-333.
Sheviak, C]. 1990. Rhodora 92: 213-231.

Spiranthes diluvialis Sheviak

UTE LADIES'- TRESSES
Arf, A. M. 1994. Aquilegia 18(2): 1, 4-5.
- 1995. NA Native Orchid Journal 1 (2): 117-128.
Sheviak, C.]. 1984. Brittonia 36: 8-14.

Spiranthes. floridana (Wherry) Cory

FLORIDA LADIES'-TRESSES

Spiranthes infernalis Sheviak

ASH MEADOWS LADIES'-TRESSES
Sheviak, C.]. 1989. Rhodora 91: 225-234.

Spiranthes magnicamporum Sheviak GREAT

PLAINS LADIES'- TRESSES

Spiranthes parksii Correll NAVASOTA

LADIES'-TRESSES
Catling, P.M. & K. L. McIntosh. 1979. SIDA 8: 188193.

The next series of examples would be those taxa that have been
described at various levels and are best treated as varieties.

Calopogon tuberosus (Linnaeus) Britton, Stems & Poggenberg

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 Brown: COLOR, FORM, AND VARIATION

var. simpsonii (Small) Magrath

SIMPSON'S GRASS-PINK

Calypso bulbosa (Linnaeus) Oakes var. americana (R.Brown)

Luer
EASTERN FAIRY-SLIPPER
Calypso bulbosa (Linnaeus) Oakes var. occidentalis (Holtzman)
Boivin
WESTERN FAIRY-SLIPPER

Coeloglossum viride (Linnaeus) Hartman var. virescens

(Mühlenberg) Luer
LONG BRACTED GREEN ORCHIS

Corallorhiza maculata (Rafinesque) Rafinesque var. occidental is

(Lindley) Ames
WESTERN SPOTTED CORALROOT
Freudenstein, J.V. 1986. Contr. Univ. Mich. Herb. 16: 145153.
- 1997. Harvard Papers in Botany) 10:5-51.

Corallorhiza odontorhiza (Willdenow) Nuttall var. pringlei

(Greenman) Freudenstein
PRINGLE'S AUTUMN CORALROOT

352
 Brown: COLOR, FORM, AND VARIATION

Freudenstein, J .V. 1993. Dissertation. Cornell

University.
- 1997. Harvard Papers in Botany) 10:5-51.

Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O.

Williams
Synonym: Corallorhiza striata forma fulva Fernald
VREELAND'S STRIPED CORALROOT

Cypripedium parviflorum Salis bury var. parviflorum Synonym:

Cypripedium calceolus Linnaeus var. parviflorum Salisbury
SOUTHERN SMALL YELLOW LADY'S-SLIPPER Sheviak,
C.J.1994. AOS Bulletin 63(6): 664-669.
- 1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal2 (4): 319-343.
Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak
Synonym: Cypripedium calceolus Linnaeus var. parviflorum
Salisbury
NORTHERN SMALL YELLOW LADY'S-
SLIPPER
Sheviak, C.J.1993. AOS Bulletin 62(4): 403.
- 1994. AOS Bulletin 63(6): 664-669.
- 1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium parviflorum Salisbury var. pubescens
(Willdenow) Knight
Synonym: Cypripedium calceolus Linnaeus var. pubescens
(Willdenow) Correll

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LARGE YELLOW LADY'S-SLIPPER

(including var. planipetalum Fernald)
Sheviak, C.J.1994. AOS Bulletin 63(6): 664-669. -
1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal 2(4): 319-343.

Cypripedium passerinum Richmond var. minganense Victorin

MINGAN SPARROW'S EGG LADY'S-SLIPPER Victorin, M.
1929. Trans. Royal Soc. Can. III 22(5): 168, pL 1-3.

Hexalectris spicata (Walter) Barnhardt var. arizonica (S. Watson)

Catling & Engel
ARIZONA CRESTED CORALROOT
Catling, P.M. & V.S. Engel 1993. Lindleyana 8(3): 119126.

Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg)

Hulten
WESTERN HEART-LEAVED TW A YBLADE

Platanthera blephariglottis (Willdenow) Lindley var. conspicua

(Nash) Luer
SOUTHERN WHITE FRINGED ORCHIS

Platanthera clavellata (Michaux) Luer var. ophioglossoides

(Fernald) P.M. Brown
NORTHERN CLUB-SPUR ORCHIS Brown,
P.M. 1988. Wild Flower Notes 3(1): 21.

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Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso)

Ledebour
BOG CANDLES
Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley)
Luer
SIERRA REIN-ORCHID

Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown)

Luer
NORTHERN TUBERCLED ORCHIS

Platanthera hyperborea (Linnaeus) Lindley var. gracilis (Lindley)

Luer
LAXLY FLOWERED BOG ORCHIS
Platanthera hyperborea (Linnaeus) Lindley var. viridiflora
(Chamisso) Luer
Synonym: Platanthera convallariifolia (Fischer) Lindley
TALL ALASKA GREEN ORCHIS

Platanthera sparsiflora (S. Watson) Schlecter var. ensifolia

(Rydberg) Luer
NARROW-LEAVED REIN-ORCHIS

Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer

Synonym: Spiranthes gracilis Bigelow
SOUTHERN SLENDERLADIES'-TRESSES

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There are a few examples of taxa that have been described

as varieties or subspecies and never reduced to synonymy.

Piperia elegans subsp. decurtata Morgan & Glicenstein PT.

REYES PIPERIA
Morgan, R & L. Glicenstein. 1993. Lindleyana 8(2): 8995.

Spiranthes casei Catling & Cruise var. novaescotiae Catling

CASE'S NOVA SCOTIAN LADIES'-TRESSES Catling,
P.M. 1981. Can. J. Bot. 59: 1253-1270

Spiranthes ovalis Lindley var. erostellata Catling

NORTHERN OVAL LADIES'-TRESSES
Catling, P.M. 1983. Brittonia 35: 120-125

The next group would be those taxa which have been

described as varieties, subspecies, or in a few cases, as
species that are best treated as forms. The are either
randomly occurring individuals in colors other than the
typical color of the species (as in white-flowered forms,
multiple-leaved forms, etc.) or environmentally induced
dwarf forms.

Arethusa bulbosa Linnaeus

DRAGON'S-MOUTH
forma albiflora Rand & Redfield - white-flowered form
subcaerulea Rand & Redfield -lilac-blue flowered

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Corallorhiza maculata (Rafinesque) Rafinesque var.

occidentalis (Lindley) Ames
WESTERN SPOTTED CORALROOT
forma aurea P.M. Brown - golden yellow/spotted form
immaculata (peck) Howell - yellow spotless form
intermedia Farwell- brown-stemmed form punicea
(Barth.) Weatherby & Adams - red- stemmed form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 195.

Galearis spectabilis (Linnaeus) Rafinesque

SHOWY ORCHIS
forma gordinierii (House) Whiting & Catling - white
flowered form
willeyi (Seymour) P.M. Brown - pink-flowered form
Brown, P.M. 1988. Wild Flower Notes 3(1): 20.

Malaxis brachypoda (Gray) Fernald

Synonym: Malaxis monophyllos (Linnaeus) Swartz. var.
brachypoda (A. Gray) Morris & Eames
WHITE ADDER'S-MOUTH
forma bifolia (Mousley) Fernald - two-leaved form

Platanthera grandiflora (Bigelow) Lindley

LARGE PURPLE FRINGED ORCHIS

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forma albiflora (Rand & Redfield) Catling - white

flowered form
bicolor P.M. Brown - bicolor-flowered form carnea
P.M. Brown - pink-flowered form mentotonsa
(Fernald) P.M. Brown - entire-lip
form
Brown, P.M. 1988. Wild Flower Notes 3(1): 22. Brown, .M.
1995. NA Native Orchid Journal 1 (1): 12. Stoutamire, W.P.
1974. Brittonia 26: 42-58.

Sacoila lanceolata (Aublet) Garay var. lanceolata

Synonym: Spiranthes lanceolata (Aublet) Leon
Spiranthes orchioides (Swartz) A. Richard
Stenorrhynchos lanceolatum (Aublet) Richard ex
Sprengel
LEAFLESS BEAKED ORCHID
forma albidaviridis Catling & Sheviak - white! green
flowered form
Catling, P. M. & C. J. Sheviak. 1993. Lindleyana. 8(2): 7781.

Triphora trianthophora (Swartz) Rydberg var. trianthophora

THREE BIRD'S ORCHIS; NODDING POGONIA forma
albidoflava Keenan - white-flowered form Keenan, P. 1992.
Rhodora 94: 38-39.

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The final group would be those taxa that represent hybrids.

Upon occasion hybrids have been described as varieties or even
species. As techniques have advances we now are able to
determine the validity of many of the putative hybrids and their
correct status.

A few examples are:

Cypripedium xalaskanum P.M. Brown ALASKAN

SPOTTED LADY'S-SLIPPER (c. guttatum x C.
yatabeanum)
Brown, P.M. 1995. NA Native Orchid Journal 1 (3): 199.

Cypripedium xandrewsii Fuller nm. andrewsii

ANDREWS'LADY'S-SLIPPER
(C candidum x C. parviflorum var. makasin)

Cypripedium xandrewsii Fuller nm. favillianum (Curtis)

Boivin
FA VILLE'S LADY'S-SLIPPER
(c. candidum x C. parviflorum var. pubescens)

Cypripedium xandrewsii nm. landonii (Garay) Boivin

LANDON'S LADY'S-SLIPPER
(C candidum x C. xandrewsii nm. favillianum)

Cypripedium xcolumbianum Sheviak

COLUMBIA LADY'S-SLIPPER (C
parviflorum x C montanum)
Sheviak, C.J.1992. AOS Bulletin 61(6): 546-559.

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Platanthera xandrewsii (Niles) Luer

Synonym: Platanthera lacera var. terrae-novae (Fernald) Luer
ANDREWS' FRINGED ORCHIS
(P. lacera x P. psycodes)
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.

Platanthera xbicolor (Rafinesque) Luer

BICOLOR FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. ciliaris)

Platanthera xcanbyi (Ames) Luer

CANBY'S FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. cristata)

Platanthera xchannellii Folsom

CHANNELL'S FRINGED ORCHIS (P.
ciliaris x P. cristata)
Folsom, J.P. 1984. Orquidea (Mex) 9(2): 344.

Platanthera xcorrellii Schrenck

CORRELL'S REIN ORCHIS (P.
hyperborea x P. stricta)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263.
Schrenck, W.J. 1978. AOS Bulletin. 47(5): 429-437.

Platanthera xestesii Schrenck

ESTES REIN ORCHIS
(P. dilatata var. albiflora x P. stricta)

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Schrenck, W.J. 1975. Die Orchidee. 26: 258-263. Schrenck,

W.J. 1978. AOS Bulletin. 47(5): 429-437.

Platanthera xkeenanii P.M. Brown

KEENAN'S FRINGED ORCHIS (P.
grandiflora x P. lacera)
Brown, P.M. 1993. A Field Guide to the Orchids of N.E. &
N.¥: p. 189.
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.

Platanthera xlassenii Schrenk LASSEN

REIN ORCHIS (P . leucostachys x
P. sparsiflora)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263. Schrenck,
W.J. 1978. AOS Bulletin. 47(5): 429-437.

Platanthera xmedia (Rydberg) Luer

INTERMEDIATE REIN ORCHIS (P.
hyperborea x P. dilatata)

Platanthera xvossii Case

VOSS' REIN ORCHIS
(P. blephariglottis var. blephariglottis x P. clavellata var.
ophioglossoides)
Case, F. W. 1983. Michigan Botanist. 22: 141-144.

Spiranthes xborealis P.M. Brown

NORTHERN HYBRID LADIES'-TRESSES (S.
casei vat. casei x S. ochroleuca)
Brown, P.M. 1995. NA Native Orchid Journal 1(4): 290.

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Spiranthes xintermedia Ames

INTERMEDIATE HYBRID LADIES'-TRESSES (S.
lacera var. gracilis x S. vernalis)
Catling, P.M. 1978. Rhodora 80: 377-389.

Spiranthes xsimpsonii Catling & Sheviak

SIMPSON'S LADIES'- TRESSES
(S. lacera var. lacera x S. romanzoffiana)
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana. 8(2): 7880.

Examples cited are often only a few of the many taxa that
would qualify under this topic. In addition there are many
undescribed colors, variations and forms that exist in the orchid
orchid throughout North America.

Paul Martin Brown, Research Associate, University of Florida

Herbarium, Gainesville, Florida. Paul is the editor of, and
frequent contributor to, this Journal.

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