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 Number 4

CONTENTS NOTES FROM THE EDITOR
297

December 1998

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

4 ANNUAL NORTH AMERICAN NATIVE ORCHID CONFERENCE
364

th

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

297

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

298

From & Read: STRATEGIES FOR CONSERVATION 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

299

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

300

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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).

301

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

302

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

303

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

Plant Tissue Histological Study Little information is available about the basic biology or culture of P. praeclara. One concern

Platanthera praeclara

304

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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 semithin 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

305

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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 humanassisted 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.

306

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

307

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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 crosspollinated 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

308

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

309

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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 noncontiguous 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.

310

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

311

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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

312

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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. 309326. Federal Register. 1989. Rules and regulations. 54: 160167. Murashige, Toshio. 1974. Plant propagation through tissue cultures. Ann. Rev. Plant Physiol. 25: 13566

313

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

314

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

315

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

316

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

317

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

318

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

319

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

320

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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).

321

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.8 P. praeclara root surface. The surfaces show fungal hyphae infection after 6 weeks in symbiotic culture. SEM at 573x.

322

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

323

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.10 Infected root cells containing bodies resembling starch grains. SEM at 1490x. When scanned individually, they indicate soft tissue.

324

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.11 Longitudinal section taken of root infected cells in symbiotic culture. SEM at 225x.

325

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.12 Cross section of the orchid root with fungal hyphae inter-connecting individual root cells. 573x SEM.

326

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.13 Infected orchid root and rhizoids in crosssection at 252x SEM. Rhizoids are single-celled extensions of epidermal hairs. They appear as hollow tubes.

327

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

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.

328

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Fig. 1.15 Tropical orchid seed which exhibits openings in the testa. The seeds of this species germinate easily in asymbiotic culture. 1640X SEM

329

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

330

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

331

From & Read: STRATEGIES FOR CONSERVATION AND PROPAGATION

Margaret M. From and Paul Read
Horticulture Department University of Nebraska Lincoln Lincoln, NE USA Omaha Henry Doorly Zoo Omaha, NE USA

332

Freeman: CYPRlPEDIUM HYBRIDS IN MAHNOMEN CO., MN

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

333

Freeman: CYPRlPEDIUM HYBRIDS IN MAHNOMEN CO., MN

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.

334

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

335

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

337

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'sslipper? 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

343

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): 171188. P Platanthera purpurascens (Rydberg) Sheviak & Jennings SHORTSPURRED BOG ORCHIS Synonym: Platanthera hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Luer Sheviak & Jennings. 1997" NA Native Orchid Journal 3(4): 444449.

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): 106116. 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): 89Piperia 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

351

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'SSLIPPER 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

353

Brown: COLOR, FORM, AND VARIATION

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.

354

Brown: COLOR, FORM, AND VARIATION

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

355

Brown: COLOR, FORM, AND VARIATION

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

356

Brown: COLOR, FORM, AND VARIATION

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

357

Brown: COLOR, FORM, AND VARIATION

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.

358

Brown: COLOR, FORM, AND VARIATION

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.

359

Brown: COLOR, FORM, AND VARIATION

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)

360

Brown: COLOR, FORM, AND VARIATION

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 hyperborea x P. dilatata)

(P.

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.

361

Brown: COLOR, FORM, AND VARIATION

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

(S.

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.

362

Brown: COLOR, FORM, AND VARIATION

363

Brown: COLOR, FORM, AND VARIATION

364

Brown: COLOR, FORM, AND VARIATION

365

Brown: COLOR, FORM, AND VARIATION

366

Brown: COLOR, FORM, AND VARIATION

367

Brown: COLOR, FORM, AND VARIATION

368