December 1998 North American Native Orchid Journal

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NORTH AMERICAN NATIVE ORCHID JOURNAL

____ __ ________________________________

Volume 4 DecemberNumber 4 1998

a quarterly devoted to the orchids of North Americapublished 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!




(ISSN 1084-7332)published quarterly in

March June September December

by theNORTH 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 scientificand general interest nature relating to the orchids of North America. Scientific articles should conform to guidelines such asthose in Lindleyana or Rhodora. General interest articles and notesmay be more informal. Authors may include line drawings,and/or black and white photographs. Color inserts may bearranged. Please send all inquiries or material for publication tothe 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 forUnited States addresses, $29US in Canada and $32US otherforeign countries. Payment should be sent to Nancy A. Webb, 84Etna St. Brighton, MA 02135-2830 USA. Claims for lost issues orcancelled memberships should be made within 30 days.




Volume 4 DecemberNumber 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 MAHNOMENCOUNTY, MINNESOTA

Rob Freeman

333

AT A LOSS?The Slow Empiricist

336

COLOR, FORM AND VARIATIONPaul Martin Brown



343

4th

ANNUAL NORTH AMERICAN NATIVE

ORCHID CONFERENCE364

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 articlesmay 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

[email protected]

Telephone & fax: 352/861-2565

mailto:[email protected]




PROCEEDINGS OF THE 3RD ANNUAL

NORTH AMERICAN NATIVE ORCHID

CONFERENCE

8-11 JULY, 1998 LAKE ITASCA,

MINNESOTAPart 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



From & Read : STRATEGIES FOR CONSERVATION AND PROPAGATION

299

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, isafforded protection under federal and state EndangeredSpecies acts. Federal and state permits were obtained toconduct this research. The species has resisted attemptsat propagation when using traditional propagationmethods, according to the Nebraska Game and ParksCommission (Fritz, 1993). Integrated conservationstrategies combining micropropagation techniques,histological studies, and in-situ species management arebeing employed for P. praeclara , whose populationnumbers are believed to be in decline. Several issues arebeing investigated in this ongoing study: 1) protocols forgerminating P. praeclara seeds in-vitro, 2) the study of seed




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structures and their possible effects on germinationresponses in-vitro, 3) histological studies of in-vitro produced P. praeclara tissues using scanning electron

microscopy and conventional high magnificationtechniques and, 4) a limited hand pollination study toassess the possibility of inducing greater fruit set within a

wild population.

Seed Germination Response

Mature seeds collected in late summer of years 1995 and1996 were air dried for five days in the lab, removedfrom the fruits and placed in sealed glass vials under

refrigeration until they were cultured. Seeds weresurface sterilized in one of two solutions; an 8% calciumhypochlorite 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 (Stoutamire1981). Frequently this occurs earliest at the suspensorend of the seed during the surface sterilization process.

This could be a factor in the damage caused to that polarregion when the sterilization solution is too harsh orapplied for too long. The carapace surrounding theembryo cells also appears dark and dense, with no visible




301

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

Germination responses were obtained on amodified Fast medium (Fast 1982), with additionalmodifications (Anderson 1990), a 1/3 strength MS(Murashige and Skoog 1962) as recommended (Chu andMudge 1994), amended with 40ml. L coconut water and6 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 coldstratification resulted in a lower germination responsethan either a 60 or 90 day cold treatment. After the coldtreatments, cultures were returned to laboratory roomtemperatures, still in continual darkness, untilprotocorms developed a root initial and a shoot initial.Protocorms which developed shoots 2 mm in height orgreater, were placed under cool white fluorescent lightsfor the remainder of the natural growing season.

Initial germination began after 150 days with 1995seeds and after as little as 18 days with 1996 seeds.However, responses were highly variable betweenindividual cultures and only a few cultures displayed 18day germination response. A small number germinatedearly, but in-vitro sown P. praeclara seeds do not easily synchronize, and germination continued for up to 17months 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 NikonFX-35 DX camera mounted on a Nikon Labophot-2.

Seeds were also sputter coated with gold andphotographed using scanning electron microscopy (SEM), on a Philips 515. Elemental analysis wasperformed with a Kevex 7000.

Dry mature seeds display a dark, opaque testa, with a distinctive, reticulate pattern unique to thespecies. The testa provides a primary barrier to wateruptake necessary for the embryo's cell division during germination. The enclosed bare embryo is surrounded

by a well-developed, dark carapace which ishypothesized to provide a secondary barrier to wateruptake, (figure 1). No visible opening is apparent in thecarapace, unlike seeds of the European native orchid,Orchis morio, whose seeds are reportedly easy togerminate in-vitro (Ronse, 1989). Seeds soaked in bleachsolution were photographed over a 2 ½ hour period torecord changes in the testa or embryos. Soaked seedsdisplayed a slightly more transparent seedcoat, which insome instances showed liquid collecting at a kevelianborder momentarily, and then rapidly sheathing thespace between and collecting at another border, (figure2). As the duration of soaking time increased, the liquidappeared to flow past the embryo with progressively greater ease. Repeated experimentation is necessary to






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frequently voiced by biologists and conservationists isthat any species propagated for reintroduction effortsmust be true to the wild genotype. Consequently, in-vitro

produced plant tissues are being photographed for plantdevelopmental studies. When the same P. praeclara seedsemi-thin section, noted above, is photographed on aconventional microscope, a small arc of yellow-gold cellsare visible at the suspensor end of the embryo. It ishypothesized that this region is the area of most rapidcell division during germination. (Appendix I, figure 1.4)

Figure 1.5, Appendix I, is a semi-thin sectionshowing considerable cell division by the time the

protocorm reaches 2 mm in length. Meristermaticregions contain the densest concentration of cells.Individual protocorm cells at 1930X SEM reveal clustersof bodies which are hypothesized to consist of mobilizedproteins and starches, (Rasmussen, 1995). Thesefracture when scanned, leading one to conclude thatthese 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 protocormsfrom 1995 began to develop a shoot initial, a root initial




306

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 rhizoidsprotruding from the surface of the spherical protocorm.Protocorms which display exceptional vigor can develop2 to 4 roots up to 12cm in length by the end of 15months. Tubers develop into a variety of shapes in-vitro,

which closely resemble those found on plants excavatedat the Sheyenne National Grasslands, (Wolken 1995).

Tubers may be coiled, rod-shaped, oval or tapered,(Appendix I fig. 1.7). Alternating cold treatments withroom temperature regimes annually appears to have a

favorable impact on tuber formation, particularly withthose protocorms which had at least one shoot 5-20mmin height. Protocorms not given an annual coldtreatment frequently become necrotic after 12continuous months at room temperature.

Hand Pollination Study

Two orchid sites were chosen to conduct human-assisted pollination. Site #1 is an upland site on aprivately-owned prairie and site #2 is a wet, low-lying prairie swale on federal lands. The sites are locatedapproximately 350 miles apart.




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The study was first conducted in 1996. Itsprimary purpose is to assess whether there is any meritin using human assisted pollination to improve seed set

for the orchid genotype(s) in wild populations located inthe far western reaches of the orchid's range, where it'spopulations are small, under threat of encroachment andthe orchid population numbers are believed to be indecline.

Twenty-eight plants were randomly chosen for study atsite #1 and 10 plants were randomly chosen at site #2.Each individual inflorescence displayed a minimum of 5fully- expanded, intact flowers. At the upland site; site#1, eight plants were cross pollinated with another

individual plant a minimum of 20 meters away. Fourplants were self-pollinated, and seven plants receiving nohuman assisted pollination were treated as a controlgroup. All plants in the study were taggedinconspicuously, staked and recorded. At the prairieswale site; site #2, located on federal lands, permission

was obtained to hand pollinate five plants and fiveadditional plants were marked as control plants.

Pollen sacs were removed from one individualflower with the aid of toothpicks and placed on thestigma region of another flower. Manipulations andstaking were completed at site #1 on June 28th, 1996.

Anthesis commenced 19 days later at site #2, and handpollination at that location took place on July 17, 1996.




308

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 perplant, at site #1. Fruit set on cross pollinated plants 5.3per plant, versus fruit set on control plants 2.7 per plant,followed a similar pattern at site #2 in 1996. Due to theextensive logistical and legal limitations placed on theexperiment at each of the two sites in 1996, the greatly reduced numbers of flowering individuals available atsite #1 in 1997, and the adverse weather conditions atsite #2 in the 1997 growing season, data are presented

here as a starting point for research in the area of humanassisted pollination to increase fruit production. Moreresearch is needed before any recommendations shouldbe made whether this practice is beneficial for long termspecies conservation. Preliminary data indicate thathuman assisted pollination may indeed increase fruit setin a wild population. Caution must be exercised sinceseed production is a costly function in terms of mostspecies' total energy reserves. Plants which were cross-pollinated also remained in a vegetative state much laterinto the autumn, and it is currently unclear whetherdelayed dormancy may affect an individual plant'slongevity. Moe and Pleasants in 1993 also questioned

whether the recorded fruit set levels they studied were




309

adequate to maintain the species at a given site. Furtherresearch is needed to adequately answer these concerns.

Assessment is needed of the threat of extinction tocurrent populations of the Western Prairie FringedOrchid. Future research into its propagation forpossible reintroduction efforts should recognize severalimportant factors: 1) the possibility that a single threatcould be capable of causing 100% mortality in asufficient 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 naturalfluctuations in populations to synchronize at a low pointthat 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 tobe available. These issues are particularly important inareas where intensive cultivation practices and theiraccompanying heavy uses of herbicides, pesticides andfertilizers, or heavy grazing practices and repeatedmowings are carried out, each of which can dramatically effect WPFO fruit-set.

Although simultaneous floods, fires or diseasepose a relatively small threat to all populationsthroughout 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' totalgenetic diversity. It is currently unknown how muchgenetic exchange occurs by natural pollen vectors among the individual sites, and future research may point to thepossibility that manipulated crossings betweenpopulations 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 northeasternportions of its range. However, this may be of littleconsequence in the western and southern portions of itsrange if individual populations do not remain viable overthe long term, and diminished populations no longerhave any genetic exchange between their disjunctlocations. Propagation as a possible source of WPFOplants in the future, may become increasingly importantfor the species' preservation and continued presence inthe western portions of the orchid's range.




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Acknowledgments

Partial funding was provided by:

Nebraska Environmental TrustIowa Living Heritage Roadway TrustNebraska Statewide ArboretumMid America Orchid Conference

Association of Zoological HorticultureOmaha Henry Doorly Zoo

Gratitude is extended to the following for theirtechnical or logistical assistance: UNL Institute of

Agricultural and Natural Resources, Marty Cano at the

University of Nebraska Medical Center in Omaha forSEM technical assistance, and Len McDaniel for fieldassistance from the U.S. Fish and Wildlife Service. Dr.Lee Simmons, Director and Terri Gouveia, HorticultureCurator at Omaha's Henry Doorly Zoo providedinvaluable assistance. Virginia Miller, UNL technologist,and the Nebraska Game and Parks Commissioncontributed technical and field insights. Karen Delaney provided histological assistance at UNMC. Drs. Karen

Johnson of University of Manitoba's Museum of Manand Nature, Devonian Botanic Garden, University of

Alberta, and Marlin Bowles at the Morton Arboretumprovided 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 andencouragement.




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Literature Cited

Anderson, A.B. 1990. Asymbiotic germination of seedsof some North American orchids, in C. E. Sayers{ed.} North American native terrestrial orchidpropagation and production. BrandywineConservancy, 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 - symbioticand 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 throughtissue cultures. Ann. Rev. Plant Physiol. 25: 135-66




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Pleasants, J. And S. Moe. 1993. Floral dispay size andpollination of the western prairie fringed orchid,

Platanthera praeclara (Orchidaceae). Lindleyana 8(1): 32-38.

Rasmussen, H. 1995. Terrestrial Orchids from Seed toMycotrophic Plant. Cambridge Press, U. K. pp.28-34

Ronse, A. 1989. In vitro propagation of orchids andnature conservation: possibilities and limitations.Mem. Soc. Roy. Belg. 11: 107-113

Stoutamire, W. 1981. Early growth in North Americanterrestrial 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 praeclaraphotographed by scanning electron microscopy (SEM).

The testa displays the characteristic reticulate patternunique to this species. Seeds of terrestrial orchids eachhave 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 browncells of the testa remnants do not stain. Photographedat 140x on a Nikon Labophot FX35.




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Fig. 1.3 (a) P. praeclara seed photographed in alongitudinal semi-thin section using SEM at 312x. Theindividual cell walls within the embryo are visible (1) andfragments 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 thelocation of those bodies in (b), within one seed cell, at625x. 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 embryoitself but none are visible in the area between theembryo and the testa.






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Fig. 1.5 (a) A semi-thin section of a P. Praeclaraprotocrom by SEM at 30X. Cell density is greatest at theapical meristem end (1). Nuclear volume has increasedas the cells enlarge and divide. Cell density at (2) may by the earliest signs of another stele or a root meristem inan early developmental stage. (b) The same protocorminterior 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 bodiesindicates that they consist primarily of potassium andcalcium. Calcium is a component of cell walls, and theanalysis may be picking up the surrounding walls.




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Fig. 1.6 The protocorm semi-thin sectionphotographed at 200x after staining with H&E.Hematoxylin stained the nuclei black (1) and all otherliving 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. 1.9 A cross-section of root cells which had notmade contact with mycelium inoculated onto thesubstrate. Individual cell walls are visible. SEM 680x.




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Fig. 1.10 Infected root cells containing bodiesresembling starch grains. SEM at 1490x. When scannedindividually, they indicate soft tissue.




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




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






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Fig. 1.14 Cross-section of infected root at 60x SEM.Note the congested appearance of root cells which hadmade contact with the mycelium in the symbioticculture.




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Fig. 1.15 Tropical orchid seed which exhibits openings in

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




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331




332

Margaret M. From and Paul Read Horticulture Department

University of Nebraska Lincoln

Lincoln, NE USA

Omaha Henry Doorly Zoo

Omaha, NE USA



Freeman: CYPRlPEDIUM HYBRIDS IN MAHNOMEN CO., MN

333

CYPRIPEDIUM HYBRIDS INMAHNOMEN 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 northernsmall 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




334

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 behaviorof 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 moreprominent hybrid features I noticed, there were many

more to numerous to mention.




335

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

[email protected]





Brown: COLOR, FORM, AND VARIATION

336

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 thatmagnifies 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




337

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 toleave 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, evensome 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




338

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 mediumbrown 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 rightterritory. 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




339

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




340

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 sphagnumand 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




341

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 theTipularia 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




342

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




343

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






345

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 AmesMRS. BRITTON'S SHADOW-WITCH

Synonym: Ponthieva racemosa (Walter) C. Mohr var. brittonae

(Ames) LuerMcCartney, 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 & LoveReinhammar, L. 1995. Nordic Journal of Botany 15(5): 469481.

- 1997. NA Native Orchid Journal 3(4): 407-425.




346

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.




347

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.




348

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.




349

Above:

Corallorhiza

striata var.

vreelandii Marin Co., CA

Right:

Cypripedium

reginae forma

albolabium

Orange Co., VT

P.M. Brown




350

Above:

Malaxis brachypoda

forma bifolia Windsor Co., VT

Right:

Epidendrum floridense

Collier Co., FL

P.M. Brown






352

var. simpsonii (Small) Magrath

SIMPSON'S GRASS-PINK

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

LuerEASTERN 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




353

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.


Cypripedium parviflorum Salisbury var. pubescens(Willdenow) Knight

Synonym: Cypripedium calceolus Linnaeus var. pubescens

(Willdenow) Correll




354

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.


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.




355

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) LuerSynonym: Spiranthes gracilis Bigelow

SOUTHERN SLENDERLADIES'-TRESSES






357

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


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




358

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-lipform

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 formCatling, 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.






360

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)




361

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)





362

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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Documents

December 1998 North American Native Orchid Journal