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!
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NORTH AMERICAN NATIVE ORCHID JOURNAL
(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.
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NORTH AMERICAN NATIVE ORCHID JOURNAL
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
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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
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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
Telephone & fax: 352/861-2565
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PROCEEDINGS OF THE 3RD ANNUAL
NORTH AMERICAN NATIVE ORCHID
CONFERENCE
8-11 JULY, 1998 LAKE ITASCA,
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
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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
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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
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and a structure resembling a small tuber. However,tuber-like structures in the first year of aseptic culture,appear to be the exception rather than the rule. The
majority of protocorms develop hair-like 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.
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Data were gathered at site #1 on September 13,1996, and results indicated a much higher average fruit
set on plants which were cross-pollinated, 5.1 per plant,than on selfed plants, 2 per plant, or control plants, 1 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
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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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Margaret M. From and Paul Read Horticulture Department
University of Nebraska Lincoln
Lincoln, NE USA
Omaha Henry Doorly Zoo
Omaha, NE USA
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CYPRIPEDIUM HYBRIDS 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
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flowering plants only, I'm quite positive there are just as many
non-flowering plants if not more.
The area is predominantly open prairie with aspen
(Populus tremuloides) wooded islands forming around prairie
potholes. The transition zone between prairie and woodland
consists primarily of Silver Buffaloberry (Shepherdia argentea). It is
within this transition zone that the majority of exotic hybrids
occur. I have seen plants with a yellow lip that was almost
completely blotched out in red, much like that of spotted
lady's-slipper, Cypripedium guttatum. On several occurrences the
lips were grotesquely deformed, possibly by a late frost.
The whole floral structure on most of the species within
this zone were also not much bigger than a dime.
Another major difference that I noticed was the 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.
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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
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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
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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
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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
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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
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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
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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
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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
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COLOR, FORM AND VARIATION
Paul Martin Brown
When does a pink lady's-slipper not look like a pink lady's-
slipper? When it has white flowers or two pouches! Many of
our native orchids can be highly variable in both color and
form. Over the years many of these variations have been
recognized at several taxonomic levels - species, subspecies,
varieties and at the forma level. I wish to review many of these
variations and discuss the appropriate level of recognition.
This paper does not intent to cover generic transfers, nor
does it intended to include all of the taxa that fall under the
respective categories. References are given for most recent
treatments of the taxa.
Those species that have been recognized at both the species
and varietal levels and are deemed to be valid species.
Some examples of this level of recognition are:
Platanthera grandiflora (Bigelow) Lindley
LARGE PURPLE FRINGED ORCHIS
Synonym: Platanthera psycodes (Linnaeus) Lindley var.
grandiflora (Bigelow) A. Gray
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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.
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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.
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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.
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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.
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Above:
Corallorhiza
striata var.
vreelandii Marin Co., CA
Right:
Cypripedium
reginae forma
albolabium
Orange Co., VT
P.M. Brown
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Above:
Malaxis brachypoda
forma bifolia Windsor Co., VT
Right:
Epidendrum floridense
Collier Co., FL
P.M. Brown
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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
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Freudenstein, J .V. 1993. Dissertation. Cornell
University.
- 1997. Harvard Papers in Botany) 10:5-51.
Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O.
Williams
Synonym: Corallorhiza striata forma fulva Fernald
VREELAND'S STRIPED CORALROOT
Cypripedium parviflorum Salis bury var. parviflorum Synonym:
Cypripedium calceolus Linnaeus var. parviflorum Salisbury
SOUTHERN SMALL YELLOW LADY'S-SLIPPER Sheviak,
C.J.1994. AOS Bulletin 63(6): 664-669.
- 1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal2 (4): 319-343.Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak
Synonym: Cypripedium calceolus Linnaeus var. parviflorum
Salisbury
NORTHERN SMALL YELLOW LADY'S-
SLIPPER
Sheviak, C.J.1993. AOS Bulletin 62(4): 403.
- 1994. AOS Bulletin 63(6): 664-669.
- 1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium parviflorum Salisbury var. pubescens(Willdenow) Knight
Synonym: Cypripedium calceolus Linnaeus var. pubescens
(Willdenow) Correll
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LARGE YELLOW LADY'S-SLIPPER
(including var. planipetalum Fernald)
Sheviak, C.J.1994. AOS Bulletin 63(6): 664-669. -
1995. AOS Bulletin 64(6): 606-612.
- 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium passerinum Richmond var. minganense Victorin
MINGAN SPARROW'S EGG LADY'S-SLIPPER Victorin, M.
1929. Trans. Royal Soc. Can. III 22(5): 168, pL 1-3.
Hexalectris spicata (Walter) Barnhardt var. arizonica (S. Watson)
Catling & Engel
ARIZONA CRESTED CORALROOT
Catling, P.M. & V.S. Engel 1993. Lindleyana 8(3): 119126.
Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg)
Hulten
WESTERN HEART-LEAVED TW A YBLADE
Platanthera blephariglottis (Willdenow) Lindley var. conspicua
(Nash) Luer
SOUTHERN WHITE FRINGED ORCHIS
Platanthera clavellata (Michaux) Luer var. ophioglossoides
(Fernald) P.M. Brown
NORTHERN CLUB-SPUR ORCHIS Brown,
P.M. 1988. Wild Flower Notes 3(1): 21.
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Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso)
Ledebour
BOG CANDLES
Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley)
Luer
SIERRA REIN-ORCHID
Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown)
Luer
NORTHERN TUBERCLED ORCHIS
Platanthera hyperborea (Linnaeus) Lindley var. gracilis (Lindley)
Luer
LAXLY FLOWERED BOG ORCHIS
Platanthera hyperborea (Linnaeus) Lindley var. viridiflora
(Chamisso) Luer
Synonym: Platanthera convallariifolia (Fischer) Lindley
TALL ALASKA GREEN ORCHIS
Platanthera sparsiflora (S. Watson) Schlecter var. ensifolia
(Rydberg) Luer
NARROW-LEAVED REIN-ORCHIS
Spiranthes lacera Rafinesque var. gracilis (Bigelow) LuerSynonym: Spiranthes gracilis Bigelow
SOUTHERN SLENDERLADIES'-TRESSES
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Corallorhiza maculata (Rafinesque) Rafinesque var.
occidentalis (Lindley) Ames
WESTERN SPOTTED CORALROOT
forma aurea P.M. Brown - golden yellow/spotted form
immaculata (peck) Howell - yellow spotless form
intermedia Farwell- brown-stemmed form punicea
(Barth.) Weatherby & Adams - red- stemmed form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 195.
Galearis spectabilis (Linnaeus) Rafinesque
SHOWY ORCHIS
forma gordinierii (House) Whiting & Catling - white
flowered form
willeyi (Seymour) P.M. Brown - pink-flowered form
Brown, P.M. 1988. Wild Flower Notes 3(1): 20.
Malaxis brachypoda (Gray) Fernald
Synonym: Malaxis monophyllos (Linnaeus) Swartz. var.
brachypoda (A. Gray) Morris & Eames
WHITE ADDER'S-MOUTH
forma bifolia (Mousley) Fernald - two-leaved form
Platanthera grandiflora (Bigelow) Lindley
LARGE PURPLE FRINGED ORCHIS
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forma albiflora (Rand & Redfield) Catling - white
flowered form
bicolor P.M. Brown - bicolor-flowered form carnea
P.M. Brown - pink-flowered form mentotonsa
(Fernald) P.M. Brown - entire-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.
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Platanthera xandrewsii (Niles) Luer
Synonym: Platanthera lacera var. terrae-novae (Fernald) Luer
ANDREWS' FRINGED ORCHIS
(P. lacera x P. psycodes)
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.
Platanthera xbicolor (Rafinesque) Luer
BICOLOR FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. ciliaris)
Platanthera xcanbyi (Ames) Luer
CANBY'S FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. cristata)
Platanthera xchannellii Folsom
CHANNELL'S FRINGED ORCHIS (P.
ciliaris x P. cristata)
Folsom, J.P. 1984. Orquidea (Mex) 9(2): 344.
Platanthera xcorrellii Schrenck
CORRELL'S REIN ORCHIS (P.
hyperborea x P. stricta)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263.
Schrenck, W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xestesii Schrenck
ESTES REIN ORCHIS
(P. dilatata var. albiflora x P. stricta)
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Schrenck, W.J. 1975. Die Orchidee. 26: 258-263. Schrenck,
W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xkeenanii P.M. Brown
KEENAN'S FRINGED ORCHIS (P.
grandiflora x P. lacera)
Brown, P.M. 1993. A Field Guide to the Orchids of N.E.&
N.¥: p. 189.
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.
Platanthera xlassenii Schrenk LASSEN
REIN ORCHIS (P . leucostachys x
P. sparsiflora)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263. Schrenck,
W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xmedia (Rydberg) Luer
INTERMEDIATE REIN ORCHIS (P.
hyperborea x P. dilatata)
Platanthera xvossii Case
VOSS' REIN ORCHIS
(P. blephariglottis var. blephariglottis x P. clavellata var.
ophioglossoides)
Case, F. W. 1983. Michigan Botanist. 22: 141-144.
Spiranthes xborealis P.M. Brown
NORTHERN HYBRID LADIES'-TRESSES (S.
casei vat. casei x S. ochroleuca)
Brown, P.M. 1995. NA Native Orchid Journal 1(4): 290.
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Spiranthes xintermedia Ames
INTERMEDIATE HYBRID LADIES'-TRESSES (S.
lacera var. gracilis x S. vernalis)
Catling, P.M. 1978. Rhodora 80: 377-389.
Spiranthes xsimpsonii Catling & Sheviak
SIMPSON'S LADIES'- TRESSES
(S. lacera var. lacera x S. romanzoffiana)
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana. 8(2): 7880.
Examples cited are often only a few of the many taxa that
would qualify under this topic. In addition there are many
undescribed colors, variations and forms that exist in the orchid
orchid throughout North America.
Paul Martin Brown, Research Associate, University of Florida
Herbarium, Gainesville, Florida. Paul is the editor of, and
frequent contributor to, this Journal.
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