NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 14 (2) 2008

IN THIS ISSUE:

SYNONYMS & MISAPPLIED NAMES OF WILD ORCHIDS FOUND IN NORTH AMERICA NORTH OF MEXICO ORCHID FLORA OF THE PANTHER NATIONAL WILDLIFE REFUGE CONTRIBUTIONS TO THE STATUS AND MORPHOLOGY OF

PLATANTHERA PALLIDA and more………….

1

The North American Native Orchid Journal (ISSN 1084-7332) is a publication devoted to promoting interest and knowledge of the native orchids of North America. A limited number of the print version of each issue of the Journal are available upon request and electronic versions are available to all interested persons or institutions free of charge. The Journal welcomes articles of any nature that deal with native or introduced orchids that are found growing wild in North America, primarily north of Mexico, although articles of general interest concerning Mexican species will always be welcome.

2

NORTH AMERICAN NATIVE ORCHID JOURNAL
Volume 14 (2) 2008

CONTENTS NOTES FROM THE EDITOR 68 NEWS OF INTEREST: SPIRANTHES DILUVIALIS NEW TO CANADA 69

ORCHID FLORA OF THE FLORIDA PANTHER NATIONAL WILDLIFE REFUGE SYNONYMS AND MISAPPLIED NAMES OF WILD ORCHIDS FOUND IN NORTH AMERICA NORTH OF MEXICO A NEW COMBINATION MY FAVORITE THINGS THE SPIRANTHES FORMERLY KNOWN AS PARKSII
Lucy A. Dueck

Scott L. Stewart & Larry W. Richardson 70

105

138

Connie Bransilver 139 143

CONTRIBUTIONS TO THE STATUS AND MORPHOLOGY OF PLATANTHERA PALLIDA, PALE FRINGED ORCHIS COMING IN JULY……
Robert T. McGrath 150 157

PLATANTHERA PALLIDA -

FIFTEEN YEARS OF COMPARISONS

Paul Martin Brown 158

66

GUARD CELLS IN SPIRANTHES STAYING HEALTHY HYBRIDS IN THE GENUS CALOPOGON
Paul Martin Brown 176
NEW TAXA AND COMBINATIONS IN THIS ISSUE 181 BOOK REVIEWS & PUBLICATION ANNOUNCEMENTS

Robert J. Ferry 167

The Slow Empiricist 174

182
Unless otherwise credited, all drawings in this issue are by Stan Folsom. The opinions expressed in the Journal are those of the authors. Scientific articles may be subject to peer review and popular articles will be examined for both accuracy and scientific content. Volume 14 (2): 68-184 issued April 3, 2008. Copyright 2008 by the North American Native Orchid Journal Cover: Listera australis by Stan Folsom

67

NOTES FROM THE EDITOR This issue is packed with a wide variety of articles, both original, and reprinted from other publications. Lucy Dueck has done a masterful job of taking a highly technical paper on Spiranthes parksii she and Ken Cameron wrote for Conservation Genetics and summarizing it for a general readership as well as adding a few more points to the paper. Bob Ferry, editor of the McAllen International Orchid Society Journal has authored a very unusual paper on leaf stomata in Spiranthes originally published in March 2008 in his journal and reformatted for us here. I was recently asked to review an article for the Long Island Botanical Society Newsletter on Platanthera pallida, a species I described in 1993, and was so impressed with it I asked to reprint it here in this issue. The upshot was then a request from Eric Lamont of LIBS for me to write a fifteen year review of the species that will appear here first and then be printed simultaneously in the April LIBS Newsletter; a fine example of reciprocal publications. This is by far the largest issue of the NANOJ ever and the featured articles conclude with Scott Stewart and Larry Richardson‘s orchid flora of the Florida Panther National Wildlife Refuge in southern Florida. Paul Martin Brown Editor naorchid@aol.com 10896 SW 90th Terrace, Ocala, FL 34481 PO Box 759, Acton, Maine 04001 June- mid October

68

SPIRANTHES DILUVIALIS NEW TO CANADA
Spiranthes diluvialis, Ute ladies‘-tress, has recently been reported as a new record from both British Columbia and Canada (Bjork et al 2008, BEN #391) and is a range extension of about 20 km from the nearest population in Okanogan County, Washington State (Bjork et al. 2008; Fertig et al 2005) where it was discovered by Bjork in 1997. A globally rare species, it is listed at the federal level as Threatened in the United States where it is known from widely scattered occurrences in the western states, east to northwestern Nebraska (Fertig et al. 2005). It is distinguished from S. romanzoffiana by its spreading lateral sepals, its longer and denser hairs in the inflorescence, and its lack of a distinctly flared distal portion of the lip (Sheviak & Brown, 2002). A single plant of S. diluvialis (photos verified by Charles Sheviak) was found growing in marshes on the eastern shore of Osoyoos Lake on Osoyoos Indian Band land. Earlier, plants of Spiranthes were found at Mahoney Lake, about 3 km east of Vaseux Lake, by Ole and Greta Westby and subsequently identified by Curtis Bjork as S. diluvialis. Due to its rarity, no specimen was collected, but a photo voucher will be deposited at the University of British Columbia Herbarium. Spiranthes diluvialis grows in diverse habitats, but unlike S. romanzoffiana Cham., it does not grow in bogs or fens, and it appears to have a preference for calcareousus or moderately saline soils. In British Columbia Spiranthes diluvialis growing in British Columbia and Washington, it is associated with Carex photo by C. Bjork viridula Michx. and/or Eleocharis rostellata (Torr.) Torr. Please note that both sites are on private lands and the specific permission is required to enter those lands. This is certainly one of the rarest plants in Canada, so every individual should be guarded from damage.
REFERENCES: Botanical Electronic Newsletter # 391 March 25, 2008 http://victoria.tc.ca/mailman/listinfo/ben-l Bjork, C.R., T. McIntosh, and R. Hall. 2008. Noteworthy Collections: British Columbia. Madrono 54: 366-367. Fertig, W., R. Black and P. Wolken. 2005. Rangewide Status Review of Ute Ladies'-Tresses (Spiranthes diluvialis). http://www.fws.gov/mountain-prairie/species/plants/uteladiestress/SPDI_Status%20review_Fertig2005.pdf Sheviak, C.J. and P.M. Brown. 2002. Spiranthes in Flora of North America Editorial Committee. Flora of North America north of Mexico. Vol. 26. New York. Oxford University Press. This item of interest is based on reports in Madrono and BEN as referenced above and communications with Curtis Bjork and Terry McIntosh. 69

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

ORCHID FLORA OF THE FLORIDA PANTHER NATIONAL WILDLIFE REFUGE
Scott L. Stewart, Ph.D. & Larry W. Richardson INTRODUCTION The Orchid Flora of the Florida Panther National Wildlife Refuge project was initiated in 2000 with the start of a comprehensive orchid conservation program at the refuge. Originally intended as a short-term project to identify species for priority conservation efforts and eventual reintroductions, this flora has grown into a regularly updated and official inventory of the orchid diversity present on the Florida Panther National Wildlife Refuge (FPNWR). The FPNWR is a 26,400 acre public land located in north-central Collier County (Florida) managed by the U.S. Fish and Wildlife Service. The FPNWR is mostly closed to public access in order to best protect the Federally-endangered Florida Panther, as well as the many other rare, threatened, and endangered animals and plants known on the refuge. The refuge is part of a unique collection of protected public lands in southwestern Florida (Collier and Monroe Counties) that comprises the Big Cypress Basin eco-region—a massive wetland area of surface water collection, drainage, and aquifer recharge for the southwest portion of the Florida peninsula. Located directly north of the Fakahatchee Strand Preserve State Park, sharing an eastern boarder with Big Cypress National Preserve, and located south of Corkscrew Swamp Sanctuary, the FPNWR represents the northern portion of the greater Fakahatchee Strand swamp. During the wet season, water slowly flows from the Corkscrew Swamp and Big Cypress areas southwesterly, through the FPNWR, and into the Fakahatchee Strand State Preserve Park before entering the Gulf of Mexico through 10,000 Islands National Wildlife Refuge (sister refuge to FPNWR). Orchid hunting on the FPNWR can be a unique experience. There are no paved roads on this wildlife refuge—no need for paved roads since the FPNWR is almost entirely closed to the public. In fact, the only paved roads near the refuge make up the southern and eastern borders of this public land. Most ―roads‖ are simple dirt and rock paths just wide enough for one pickup truck, originally cut by hunters, cattle ranchers, and loggers decades ago. Many of the roads are now maintained as fire breaks. Of course, there are no direct routes to any area of the refuge, so what may appear to be a simple twenty minute walk to a particular cypress dome when looking at a map may, in fact, be a hour drive and walk to get close to the dome! Complicating matters even further, those same dirt and rock roads transform into miles and miles of boot-swallowing mud when the summer rains arrive. Fortunately, a little south Florida creativeness has provided an answer to summer orchid hunting—the swamp buggy. This truly all-terrain vehicle allows refuge personnel, biologists, and wildland fire fighters access to nearly all parts of the FPNWR no matter the conditions. Once the swamp buggy or pickup truck has taken you as far as the road will go, ambitious orchid hunters still may have a mile or so slog through knee-deep water, a long hike across an open prairie with the south Florida sun blazing overhead, or a ―walk‖
70

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

through an eerily still swamp mid-chest deep in water in order to find, catalog, and photograph the native orchids of this amazing place. And one can not forget about the nearly ever-present insects, especially those mosquitoes. In an effort to conserve the native orchid diversity present on the FPNWR, a conservation program was started in 2000. This program, initially, focused on the collection of seed from all orchid species on the FPNWR and subsequent propagation of those species; with the eventual goal of reintroducing resulting plants back into historic and restored habitats on the refuge. In 2002, the program expanded through the establishment of a partnership with Dr. Michael Kane‘s research program in the Environmental Horticulture Department at the University of Florida. Also in that same year, the FPNWR orchid conservation program began to adopt the concept of integrated orchid conservation as the primary concept behind all orchid conservation research on the refuge (Stewart, 2007; Stewart and Kane, 2007; Stewart et al., 2007). This approach combines the study of orchid ecology, mycology, propagation, pollination biology, genetic diversity, and reintroduction science with the overall goal of orchid species recovery (Hopper, 1997; Stewart, 2007). In 2005, Dr. Lawrence Zettler‘s orchid conservation program from Illinois College and his undergraduate research students joined the FPNWR orchid conservation program. The Zettler lab, thus far, has focused mostly on research with the epiphytic orchids of the refuge—dealing mainly with symbiotic and asymbiotic propagation, diversity, and basic ecology. To date, the FPNWR orchid conservation program has exercised this integrated model for a number of south Florida native orchids, including Cyrtopodium punctatum (D. Dutra, personal communication), Bletia purpurea (D. Dutra et al., personal communication), Eulophia alta (Johnson et al., 2007), Calopogon tuberosus var. tuberosus and var. simpsonii (P. Kauth, personal communication), Spiranthes odorata (S.L. Stewart, unpublished data), Dendrophylax lindenii (S.L. Stewart, unpublished data), Epidendrum nocturnum (Zettler et al., 2007), and Prosthechea cochleata var. triandra (S.L. Stewart, unpublished data; L. Zettler, personal communication). The ultimate goal of the FPNWR orchid conservation program is the systematic development of conservation and recovery plans for each of the twenty-seven currently known species present on the refuge. The Orchidaceae of the FPNWR are described herein. The last complete, orchid floristic treatment of this public land was prepared by S.L. Stewart (2003), and only provided a brief synopsis of the orchid diversity present on the FPNWR. The present treatment includes twentyseven species in seventeen genera. This flora differs substantially from the previous work on the orchid flora of the FPNWR because of nomenclatural changes, new records, the discovery of new species, the inclusion of synonymous nomenclature, and the inclusion of color and growth forms. None of the included species are strictly endemic to the FPNWR, but two species are considered narrowly restricted to the southwestern Florida counties. A number of species treated in this flora have records only in southern Florida and Cuba. NATURAL COMMUNITIES OF THE FPNWR The FPNWR is comprised of a number of diverse and orchid-rich natural communities, most of which are common throughout southwestern Florida. Unlike the expansive Everglades Sea of Grass and tropical hammock systems of southeastern Florida, southwestern Florida is dominated by seasonally inundated cypress forests and domes, sloughs and strand swamps, marl prairies, and pine forests. On the FPNWR, the predominant natural communities include hardwood hammocks, pine forests, wet prairies, cypress domes, and strand swamps and sloughs. Small lakes and ponds are also quite common throughout the FPNWR, and are typically
71

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

surrounded by deepwater sloughs. As with nearly all natural community types in southern Florida, water is the primary ecological factor—creating discrete seasons of dry and wet on the refuge. The following paragraphs present brief synopses of key characteristics of the major natural communities on the FPNWR. A diagnostic image of the natural community follows each synopsis. Hardwood Hammocks Hardwood hammocks, also termed rockland hammocks or mesic hammocks, are reasonably common on the FPNWR. They are typically comprised of a sandy limestone soil, with occasional exposed limestone, and pockets of wetter depressions. On the FPNWR, these hardwood communities are often dominated by live oak (Quercus virginiana), cabbage palm (Sabal palmetto) and sometimes gumbo limbo (Bursera simaruba) trees, with occasional saw palmetto (Serenoa repens). Fire is nearly absent from these habitats due to generally high moisture content in both the hammock soil and detritus on the hammock floor. Native orchids that can be found in this natural community include the genera Habenaria and Oeceoclades. Epiphytic orchids are occasionally found in hardwood hammocks, although this is not common on the FPNWR.

A typical hardwood hammock on the FPNWR. Slash pine (Pinus elliotti) is an occasional component of these hammocks in southwestern Florida and typically indicates a slightly dryer hardwood hammock.

Pine Flatwoods Pine flatwoods/forests, also known as mesic flatwoods, are typically dominated by slash pine (Pinus elliotti) and a mid- and understory of cabbage palm and saw palmetto. A number of assorted grasses and other low-growing herbaceous plants are common in southwestern Florida mesic pine flatwoods. Fire is common in this natural community, and is regularly used as a management tool in these communities on the FPNWR (FPNWR, 2000). Native orchids common to this natural community include the genera Bletia, Calopogon, and Eulophia. While it is uncommon to find epiphytic orchids growing in this seasonally-dry community, Encyclia tampensis can occasionally be found in pine flatwoods.

72

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

A typical pine flatwood on the FPNWR.

Wet Prairies Wet prairies, also called marl prairies, are one of the predominant natural communities on the FPNWR. This type of prairie is seasonally inundated and often is comprised of sawgrass (Cladium jamaicensis), muley grass (Muhlenbergia capillaries), maidencane (Panicum hemitomon), other assorted grasses, and occasional dwarf cypress (Taxodium distichum). Fire is common in wet prairies and is used as the primary means of management for this habitat (FPNWR, 2000). A number of terrestrial native orchids can be found in the wet prairies, including species of the genera Calopogon, Bletia, Spiranthes, and Zeuxine. Epiphytes are uncommon in this natural community given the lack of trees; however, in wet prairies where dwarf cypress trees are present Encyclia tampensis and Cyrtopodium punctatum could be present. Cypress Forests Cypress forests, also commonly called floodplain swamps, are another common natural community on the FPNWR. Cypress (Taxodium distichum) dominates this seasonally inundated natural community. The understory of most cypress forests is comprised of buttonbush (Cephalanthus occidentalis), pickerelweed (Pontederia spp.), and occasional cabbage palm, depending on the history of flooding and hydroperiod. Slightly deeper depressions within cypress forests may be dominated by pop ash (Fraxinus caroliniana) trees and associated wetland species. A special type of cypress forest, the cypress dome, is common on the FPNWR. Cypress domes are deeper depressions in the cypress forest soils where cypress trees of increasing height are found closer to the center of the depressions in response to deeper organic soils. Native orchids common to cypress forests and cypress domes include the genera Eulophia, Spiranthes, Harrisella, Encyclia, and Epidendrum.
73

A wet prairie on the FPNWR with spider lilies (Hymenocallus palmeri).

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

A cypress dome in profile (top) and cypress domes surrounded by pine flatwoods from the air (bottom).

Sloughs and Strand Swamps Strand swamps and sloughs are common natural communities on the FPNWR. Sloughs are characterized as broad, reasonably shallow wetland channels having peat soils that are seasonally inundated with water flowing slowly in response to a nearly flat topography. Sloughs are typically dominated by pop ash and pond apple (Annona glabra) trees with occasional cypress and red maple (Acer rubrum). Similar to sloughs, strand swamps are more linear wetland channels with peat soils and seasonal inundation. Again, slowly flowing water is a key characteristic of the strand swamp. Strand swamps are often deeper than sloughs, and are dominated by cypress, pop ash, pond apple, and willow (Salix caroliniana). Many native orchids can be found within both sloughs and strand swamps, including species in the genera Campylocentrum, Dendrophylax, Encyclia, Epidendrum, Ionopsis, Polystachya, and Prosthechea.

74

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Typical swamp and slough on the FPNWR. Note the floating logs, dense vegetation, and standing water.

Taxonomic Concepts The taxonomy utilized in this flora follows that of Brown (2002, 2005). Synonyms indicated may be found in Luer (1972, 1975) and the Flora of North America North of Mexico volume 26, Orchidaceae (2002). Ackerman (1995) and Llamacho and Larramendi (2005) have been used to cross-reference species accounts and synonymous nomenclature. Every attempt has been made to include appropriate and common synonyms, as well as misapplied names, where possible. Orchid Flora This section contains accounts for each species currently known from the FPNWR. Each account is arranged first by scientific name (and synonyms), then by common name, typical flowering period on FPNWR, habitat description as it pertains to their occurrence on the refuge, a brief description of the plant, and a short narrative about the species on the FPNWR. Specific location information within the FPNWR has been purposefully removed from this floristic treatment at the request of the U.S. Fish and Wildlife Service and the FPNWR.

75

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Bletia purpurea (Lamark) de Candolle Common name: pine-pink Flowering period: December to March in southeastern Florida (Everglades), April to May in the Fakahatchee/FPNWR region; can flower any time of year Habitat: Open wet pine flatwoods, wooded swamps, wet/damp roadsides, and slightly disturbed sites Description: terrestrial up to 2 m tall; 3-5 green linear lanceolate leaves; 5-20 pink to pinkpurple flowers; lip with gold/yellow crests; individual flower size 1.5-5 cm Bletia purpurea is one of the more commonly encountered orchids on the FPNWR, with formally wide distributions throughout the pine flatwoods, cypress forests, and wet prairies on the refuge. However, the species is probably now less abundant in these natural communities due to habitat changes over the past 50-plus years. While a white-flowered form, forma alba (Ariza-Julia & Jimenez Alm.) P.M. Brown, is known from sites in south Florida, this form is currently not known on the FPNWR. Additional searches are likely to identify forma alba on the refuge. Currently, an intensive research project has been undertaken by students at the University of Florida focused on better understanding the integrated conservation of B. purpurea.

Plant in capsule.

76

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Calopogon multiflorus Lindley Synonyms: Calopogon barbatus (Walter) Ames var. multiflorus (Lindley) Correll Limodorum multiflorum (Lindley) Mohr Limodorum pinetorum Small Common name: many-flowered grass pink Flowering period: (March) April (July), depending on fire management regime Habitat: damp prairies and pine flatwoods Description: terrestrial up to 30 cm tall; 1 or 2 slender leaves shorter than the height of the plant; 3-10 non-resupinate pink flowers; lip with gold/yellow crests; individual flower size 2-2.5 cm Calopogon multiflorus is one of the rarest orchids on the FPNWR, discovered several years ago in a pine flatwood approximately 4 weeks after a prescribed fire. The species has not been seen since despite the same location being exposed to prescribed fire in subsequent years. The species has likely been overlooked in other pine flatwood habitats. Future searches may also discover the white-flowered form of this species, forma albiflorus P.M. Brown.

77

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. tuberosus Synonyms: Calopogon pulchellus (Salisbury) R. Brown Calopogon pulchellus (Salisbury) R. Brown var. latifolius (St. John) Fernald Limodorum pulchellum Salisbury Limodorum tuberosum Linnaeus Common name: common grass-pink Flowering period: March to August Habitat: wet prairies, mesic pine flatwoods, and damp roadsides Description: terrestrial up to 75 cm tall; 1-4 slender leaves shorter than the height of the plant; 5-20 non-resupinate deep to pale pink flowers; lip with gold crests; individual flower size 2-3.5 cm As with Bletia purpurea, Calopogon tuberosus var. tuberosus is one of the more commonly encountered native orchids in the wet prairies and pine flatwoods on the FPNWR. The white-flowered form of this species, forma albiflorus Britton, is known from the FPNWR. Students from the University of Florida are currently studying the conservation, propagation, and ecotypic differentiation of this species.

Typical rose-pink color.

Forma albiflorus from the FPNWR.

78

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. simpsonii (Small) Magrath Common name: Simpson‘s grass-pink Flowering period: late December to June Habitat: open marl prairies Description: terrestrial up to 120 cm tall; 1-5 slender leaves typically shorter than the height of the plant; 5-10 non-resupinate pink flowers; lip with gold crests; individual flower size 2-3.5 cm Calopogon tuberosus var. simpsonii is a locally common orchid on the FPNWR. This variety grows and blooms sympatrically with var. tuberosus on the refuge, which may have resulted in intermediate hybrids between the two varieties. Plants can be easily found that exhibit intermediate characteristics of both parental varieties. Research by students at the University of Florida is attempting to describe the genetic differentiation and diversity of these sympatric populations on the FPNWR, along with pollination biology, mycological, and propagation differences.

79

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Campylocentrum pachyrrhizum (Reichenbach f.) Rolfe Common name: ribbon orchid; crooked-spur orchid Flowering period: September to November Habitat: hardwood hammocks, cypress forests, sloughs, and strand swamps Description: epiphytic; leafless; flattened gray-green roots; up to 30 pale yellow flowers on pendant stems; individual flower size 3-5 mm. Campylocentrum pachyrrhizum is known from only a few locations on the FPNWR, although it is locally abundant at the neighboring Fakahatchee Strand Preserve State Park. This species is sometimes difficult to spot when surveying its swamp and slough habitat, and is therefore likely overlooked on the refuge.

80

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Cyrtopodium punctatum (Linnaeus) Lindley Common name: cigar orchid; cowhorn orchid Flowering period: March to May Habitat: hardwood hammocks, open sunny swamps and cypress forests Description: epiphytic; pseudobulb fusiform; flowering panicle up to 1 m long; 10-15 palm-like leaves; 30-70 flowers with pale yellow-green sepals marked with reddish; petals brighter yellow with fewer markings; lip reddish with yellow and orange markings near center; individual flower size 4-6 cm Cyrtopodium punctatum is one of the most impressive orchids encountered on the FPNWR. The species is not common and is widely scattered—with only a handful of known plants on the refuge. Additional populations of the species are known from surrounding public lands. Students from the University of Florida have been studying the pollination biology, mycology, and propagation of this orchid for several years, and have developed conservation and reintroduction plans for the species. In the coming years, these conservation plans will likely result in continued reintroductions of C. punctatum onto the refuge.

81

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Dendrophylax lindenii (Lindley) Bentham ex Rolfe Synonyms: Polyradicion lindenii (Lindley) Garay Polyrrhiza lindenii (Lindley) Cogniaux Common name: ghost orchid, frog orchid Flowering period: May to August, although individuals have been known to flower any time of year Habitat: hardwood hammocks, sloughs, strand swamps, cypress domes Description: epiphytic; leafless with gray-green roots; 1-3 flowers typical; sepals and petals lanceolate; lip 3-lobed; central lobe triangular; side lobes elongated and twisting; spur slender up to 15 cm long; individual flower size to 12 cm Dendrophylax lindenii is one of the most striking orchids encountered on the FPNWR. The species is known only from a few locations deep within the interior of the refuge, although continued surveys are likely to discover additional plants and populations. While single-flowered individuals are normal, two and three-flowered individuals are also known from the refuge and surrounding conservation areas. Research efforts by students at the University of Florida are currently underway to develop conservation and reintroduction plans for this species.

Note one open flower and two buds.

82

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Encyclia tampensis (Lindley) Small Synonym: Epidendrum tampense Lindley Common name: Florida butterfly orchid Flowering period: June to July, although individuals have been known to flower throughout the year Habitat: pine flatwoods, hardwood hammocks, sloughs, strand swamps Description: epiphytic; pseudobulb elliptic to ovoid; inflorescence up to 80 cm long; 1-2 leaves; up to 30 yellow, copper, green, and/or bronze flowers; sepals and petals similar; individual flower size 3-5 cm Encyclia tampensis is one of the most commonly encountered orchids on the FPNWR. The species is locally abundant in the swamps and flatwoods throughout the refuge. The lip shape and color pattern can show great variation from plant to plant. The white-lipped form, forma albolabia (A. Hawkes) E. Christensen, while not currently known from the FPNWR is likely present.

83

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Epidendrum amphistomum A. Richard Epidendrum anceps Jacquin – misapplied Common name: dingy-flowered star orchid Flowering period: January to July, although may flower throughout the year Habitat: sloughs and strand swamps Description: epiphytic; erect stems up to 120 cm tall; 5-20 green elliptic leaves; up to 30 bronzy-green flowers; sepals and petals similar; individual flower size 1.5-2.5 cm Epidendrum amphistomum is another commonly encountered orchid in the swamps and cypress domes on the FPNWR. The red-leaved form, forma rubrifolium P.M. Brown, is known from only one location on the FPNWR.

84

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Epidendrum floridense Hágsater Epidendrum difforme Jacquin – misapplied Epidendrum umbellatum Swartz – misapplied Neolehmannia difformis (Jacquin) Pabst – misapplied Common name: Florida star orchid Flowering period: August to November Habitat: deepwater sloughs and strand swamps Description: epiphytic; pendant stems up to 30 cm long; 5-10 elliptic glossy green leaves; up to 30 yellow-green flowers; sepals and petals similar; individual flower size 1.5-3 cm Epidendrum floridense was reported from one location deep in the interior of the FPNWR several years ago. Despite two subsequent surveys, this original location or additional plants could not be located. However, the species is included in the current flora because of the reliability of the original source. Additional searches for this species are being conducted as part of the refugewide orchid conservation program.

85

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Epidendrum nocturnum Jacquin Common name: night-fragrant epidendrum Flowering period: July to January, although individuals may flower any time of year Habitat: sloughs and strand swamps, and cypress domes Description: epiphytic; erect stems to 110 cm tall; 4-10 elliptic dark green leaves; up to 3 pale yellow and white flowers; sepals yellowish and slender with petals similar; individual flower size up to 12 cm Epidendrum nocturnum is a locally abundant orchid found in the swamps and cypress forests on the FPNWR. The species appears to be self-pollinating in Florida, with multiple capsules being a common sight on large plants. Conservation research by Dr. Lawrence Zettler at Illinois College has resulted in plants being reintroduced into select sites on the refuge.

86

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Epidendrum rigidum Jacquin Common name: rigid epidendrum Flowering period: October to May Habitat: sloughs and strand swamps, and cypress domes Description: epiphytic; short, erect stems up to 20 cm tall; 2-10 elliptic green leaves; 3-10 sessile, non-resupinate green flowers; individual flower size approximately 5 mm Epidendrum rigidum is a common orchid in the cypress domes, sloughs, and strand swamps on the FPNWR. This species is occasionally found in heavily shaded cypress forests as well. Propagation and plant reintroduction research is currently being conducted by undergraduate students at Illinois College.

87

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Eulophia alta (Linnaeus) Fawcett & Rendle Common name: wild coco Flowering period: (July) August to December (January) Habitat: wooded swamps, pine flatwoods, wet prairies Description: terrestrial; 20-180 cm tall; 4-6 yellow-green leaves; up to 30 pink to green to light yellow flowers; individual flower size 3-4.5 cm Eulophia alta is a locally common orchid on the FPNWR. The species exhibits a high degree of variability in flower color, although the pale-colored form, forma pallida P.M. Brown, and the white and green-flowered form, forma pelchatii P.M. Brown, have not been observed on the refuge. Students from the University of Florida are currently conducting pollination biology and propagation research with this species that will lead to plant reintroductions.

Vegetative plants with typical palm-like leaves.

88

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Habenaria odontopetala Reichenbach f. Synonyms: Habenaria strictissima Reichenbach f. var. odontopetala (Reichenbach f.) L.O. Williams Habenella odontopetala (Reichenbach f.) Small Habenaria floribunda Lindley – misapplied Common name: toothed Habenaria Flowering period: February, in south Florida Habitat: hardwood hammocks and pine flatwoods Description: terrestrial; up to 1 m tall; 5-10 elliptic glossy green leaves; 10-50 flowers with green sepals and yellow-green petals; individual flower size 2 × 2.5 cm Habenaria odontopetala is a locally common orchid in the hardwood hammocks and roadside edges of pine flatwoods throughout the FPNWR. In pine flatwoods, the species is often found growing in pine needle duff.

89

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Habenaria quinqueseta (Michaux) Eaton Synonyms: Habenaria michauxii Nuttall Habenaria simpsonii Small Platanthera michauxii (Nuttall) Wood Common name: Michaux‘s orchid Flowering period: January, in south Florida Habitat: hardwood hammocks and pine flatwoods Description: terrestrial; up to 30 cm tall; 3-6 elliptic glossy green leaves; 8-15 flowers; sepals oblong and light green; petals white; individual flower size approximately 4 × 4 cm Habenaria quinqueseta was a surprise discovery on the FPNWR, unexpectedly found growing in a clearing in a pine flatwood along with Bletia purpurea. To date, no other plants have been found; although, additional plants are likely to occur in similar natural communities on the refuge.

90

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Habenaria repens Nuttall Synonyms: Habenaria nuttallii Small Platanthera repens (Nuttall) Wood Common name: water-spider orchid Flowering period: free-flowering throughout the year Habitat: sloughs, strand swamps, cypress domes, and cypress forests Description: terrestrial or aquatic; up to 50 cm tall; 3-8 yellow-green leaves; 10-30 light green flowers in a densely packed raceme; individual flower size approximately 2 × 2 cm Habenaria repens is one of the most common orchids throughout Florida; however, until 2006 the species was not known on the FPNWR. Several flowering plants were found on floating logs in a slough at that time, and several additional plants have been found in other sloughs and strand swamps throughout the refuge since. Dense populations are not common on the FPNWR. Additional searches should yield more populations of this common species.

91

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Harrisella porrecta (Reichenbach f.) Fawcett & Rendle Synonyms: Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten Harrisella filiformis (Swartz) Cogniaux – misapplied Common name: leafless harrisella Flowering period: August to November Habitat: hardwood hammocks, cypress domes, cypress forests, sloughs, and strand swamps Description: epiphytic; leafless; roots gray-green and slender; up to 5 slender panicles of 1-8 non-resupinate pale yellow flowers; individual flower size approximately 2.5 × 3 mm Harrisella porrecta is known from only two locations on the FPNWR—one a deepwater slough and the other a disturbed depressional area in a pine flatwood/cypress forest community. This species is likely more widely distributed on the refuge, but has been overlooked.

92

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Ionopsis utricularioides (Swartz) Lindley Common name: delicate ionopsis Flowering period: December to April Habitat: hardwood hammocks, sloughs, strand swamps, and cypress domes Description: epiphytic; pseudobulbs green and concealed within leaf bases; 1-5 olive green lanceolate leaves; 3-40 white-pink flowers with dark pink veining on a many-branched panicle; individual flower size 1 × 1.5 cm Ionopsis utricularioides is an uncommon orchid on the FPNWR, and is only known from a few locations along a tram in a mixed slough/cypress forest community. As with many of the orchids occurring on the FPNWR, additional locations for this species have likely been overlooked or never explored.

93

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Oeceoclades maculata (Lindley) Lindley Common name: spotted African orchis Flowering period: August to November Habitat: hardwood hammocks and cypress forests Description: terrestrial; pseudobulb ovoid; 1-3 dark green mottled leaves; 3-10 white flowers with purple markings; petals and sepals similar; individual flower size 1-2 cm Oeceoclades maculata is a naturalized exotic orchid in Florida and persistent populations are quite common in the southern half of the state. On the FPNWR, this species was originally only known from a few plants in one small hardwood hammock on the boarder of the refuge. However, more recent surveys have identified scattered plants and populations of O. maculata from deep within the interior of the FPNWR.

94

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Polystachya concreta (Jacquin) Garay & Sweet Polystachya flavescens (Lindley) Small – misapplied Common name: helmet orchid Flowering period: September to December, although plants may flower any time of year Habitat: hardwood hammocks, cypress forests, cypress domes, sloughs, and strand swamps Description: epiphytic; typically forms large masses; pseudobulbs cylindrical and tapered; 2-5 lanceolate-linear leaves; 10-50 pale yellow non-resupinate flowers typically on a branched raceme Polystachya concreta is a locally common orchid on the FPNWR, capable of being found in a variety of damp natural community. This species is known to occur as a single plant on a tree substrate or as large clumps of plants stretching for a meter or more. Efforts by students at Illinois College are currently underway to study the propagation and ecology of this species on the FPNWR.

95

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Synonyms: Encyclia bidentata (Lindley) Hágsater & Soto Arenas subsp. erythronioides (Small) Hágsater Encyclia boothiana (Lindley) Dressler var. erythronioides (Small) Luer Epicladium boothiana (Lindley) Small var. erythronioides (Small) Acuña Pseudencyclia boothiana (Lindley) V.P. Castro & Chiron Common name: Florida dollar orchid Flowering period: August to November Habitat: swamps and strand swamps Description: epiphytic; pseudobulbs circular and flattened; 1-3 thin oblanceolate green leaves; 2-8 yellow to tan flowers with brown or purplish markings; sepals and petals similar; individuals flower size 2-4 cm Prosthechea boothiana var. erythronioides is an extremely uncommon orchid on the FPNWR, and is currently known from only one plant in one location. This plant was discovered accidentally while exploring a deepwater slough for Roseate Spoonbill roosting sites. The plant has not been seen in flower; however, the flattened and circular pseudobulbs are diagnostic for this species

96

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins Synonyms: Anacheilium cochleatum Linnaeus var. triandrum Ames Encyclia cochleata (Linnaeus) Dressler subsp. triandra (Ames) Hágsater Common name: Florida clamshell orchid Flowering period: September to May, although individuals may flower throughout the year Habitat: sloughs and strand swamps Description: epiphytic; pseudobulbs ovoid and compressed; 1-2 thin linear-lanceolate green leaves; 3-10 non-resupinate flowers; sepals and petals similar; individual flower size 2-6 cm Prosthechea cochleata var. triandra is a locally common orchid on the FPNWR, and is known to occur in a number of sloughs and strand swamps. Scattered plants and small populations can be found in nearly every slough and strand swamp on the refuge. This species has been the subject of an intensive propagation and reintroduction program on the FPNWR, which has resulted in a number of plants being successfully reintroduced into sloughs and cypress domes throughout the refuge. Despite intensive searches, the white and yellow-flowered form, forma albidoflava P.M. Brown, has not been reported on the FPNWR.

97

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Sacoila lanceolata (Aublet) Garay var. lanceolata Synonyms: Spiranthes lanceolata (Aublet) Leon Stenorrhynchos lanceolatum (Aublet) Richard in Sprengel Stenorrhynchos orchoides (Swartz) L.C. Richard Common name: leafless beaked orchid Flowering period: April to June Habitat: pine flatwoods and prairies Description: terrestrial; 10-60 cm tall; 4-6 oblanceolate leaves absent at flowering; 10-50 red flowers in a terminal raceme; individual flower size approximately 2 cm Sacoila lanceolata var. lanceolata was known for many years from public and private lands surrounding the FPNWR, but not known from the refuge itself. Plants of this species were accidentally discovered in a clearing in a pine flatwood while surveying for Bletia purpurea sites. Additionally, one plant of the white/green-flowered form, forma albidaviridis Catling & Sheviak, was recently discovered while visiting the same B. purpurea site near where the original plants of var. lanceolata were discovered. The golden-bronze flowered form, forma folsomii P.M. Brown, is not currently known from the FPNWR.

Typical red-coral color.

Forma albidaviridis in capsule.

98

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Spiranthes longilabris Lindley Synonyms: Ibidium longilabre (Lindley) House Spiranthes brevifolia Chapman Triorchis longilabris House Common name: long-lipped ladies‘-tresses Flowering period: November to December Habitat: pine flatwoods and wet prairies Description: terrestrial; 15-50 cm tall; 3-5 linear-lanceolate leaves withered at flowering; 10-30 white flowerings in a single rank; diagnostic widely-spreading linear sepals; individual flower size 1-1.5 cm Spiranthes longilabris is considered rare in Florida, especially in southern Florida. The species is known only from one site on the FPNWR consisting of only a few plants. As with the other Spiranthes species native to the refuge, additional S. longilabris plants and populations have likely been overlooked.

99

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Spiranthes odorata (Nuttall) Lindley Synonyms: Ibidium odoratum (Nuttall) House Spiranthes cernua (Linnaeus) L.C. Richard var. odorata (Nuttall) Correll Triorchis odorata (Nuttall) Nieuwland Common name: fragrant ladies‘-tresses Flowering period: October to January Habitat: wet prairies and pine flatwoods Description: terrestrial or semi-aquatic; 2-100 cm tall; 3-5 linear-oblanceolate green leaves; 1030 white to ivory flowers in several tight ranks; individual flower size 1-2 cm Spiranthes odorata is known from several expansive populations in the wet prairies on the FPNWR. The species was apparently overlooked for many years, despite plants occurring next to several of the commonly used roadways on the refuge. Menk (2007) and S.L. Stewart (personal observation) have recently reported the presence of a distinct green-throated S. odorata type from southwestern Florida. This new color type grows sympatrically with the typical yellowthroated color type on the FPNWR. Extensive research is being conducted on the integrated conservation of this species (both lip color types) on the refuge, particularly focusing on the propagation and ecology of the species in southwestern Florida.

Typical yellow-throated color type.

Green-throated color type.

100

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Spiranthes vernalis Engelmann & Gray Synonyms: Ibidium vernale (Engelmann & Gray) House Gyrostachys vernalis Kuntze Triorchis vernalis House Common name: grass-leaved ladies‘-tresses Flowering period: January to May Habitat: prairies and pine flatwoods Description: terrestrial; 10-60 cm tall; 2-5 linear-lanceolate green leaves typically present at flowering; 10-40 ivory to white flowers in either a single or multiple ranks; individual flower size 6-9 mm Despite being one of the most commonly encountered orchids in Florida, Spiranthes vernalis, much like Sacoila lanceolata, was known from lands surrounding the FPNWR but not known, surprisingly, from the refuge itself. Plants were first discovered in power line right-of-ways near the boarders of the refuge, and additional plants were then discovered along several of the roadways within the refuge. Plants can be locally common on the FPNWR.

101

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Scientific name: Zeuxine strateumatica (Linnaeus) Schlechter Common name: lawn orchid Flowering period: (October) December to April Habitat: prairies, pine flatwoods, and disturbed sites Description: terrestrial; 4-20 cm tall; 5-10 lanceolate leaves; 5-30+ white flowers with a yellow lip; sepals and petals similar; individual flower size 6-8 mm Zeuxine strateumatica is another non-native, naturalized exotic orchid species known from the FPNWR. The species was first discovered from disturbed roadside right-of-ways adjacent to the refuge, and subsequently reported from boarder-land pine flatwoods and prairies on the FPNWR.

102

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

Species from Adjacent Lands A number of native orchid species are known from adjacent and nearby public and private lands, but not known from the FPNWR despite the presence of analogous natural communities. As part of the ongoing comprehensive orchid conservation program at the refuge, these orchids are actively being sought on the FPNWR and may be discovered in the near future. 1) Bulbophyllum pachyrhachis – hardwood hammocks, sloughs, and strand swamps 2) Calopogon barbatus – wet prairies and pine flatwoods 3) Calopogon pallidus – wet prairies and pine flatwoods 4) Cranichis mucosa – cypress domes, sloughs, and strand swamps 5) Cyclopogon cranichoides – hardwood hammocks 6) Epidendrum acunae – deepwater sloughs and strand swamps 7) Epidendrum strobiliferum – sloughs and strand swamps 8) Gymnadeniopsis nivea – wet prairies and pine flatwoods 9) Habenaria distans – hardwood hammocks 10) Lepanthopsis melanantha – sloughs and strand swamps 11) Liparis elata – hardwood hammocks and sloughs 12) Malaxis spicata – cypress forests and sloughs 13) Maxillaria crassifolia – deepwater sloughs and strand swamps 14) Maxillaria parviflora – deepwater sloughs and strand swamps 15) Oncidium floridanum – hardwood hammocks 16) Pelexia adnata – hardwood hammocks 17) Platythelys sagreana – hardwood hammocks 18) Pleurothallis gelida – sloughs and strand swamps 19) Ponthieva racemosa – cypress forests and sloughs 20) Prescottia oligantha – hardwood hammocks 21) Prosthechea pygmaea – sloughs and strand swamps 22) Pteroglossaspis ecristata – pine flatwoods and prairies 23) Sacoila lanceolata var. paludicola – hardwood hammocks and sloughs 24) Spiranthes brevilabris – prairies 25) Spiranthes laciniata – prairies and sloughs 26) Spiranthes praecox – prairies 27) Spiranthes torta – pine flatwoods 28) Spiranthes tuberosa – pine flatwoods 29) Triphora gentianoides – disturbed areas 30) Vanilla phaeantha – sloughs and strand swamps Acknowledgements Thanks are extended to the administration and staff of the Florida Panther National Wildlife Refuge for their support of this flora project and continued support for the comprehensive orchid conservation program. The assistance of Dennis Giardina (Fakahatchee Strand Preserve State Park), Philip Kauth, Timothy Johnson, Daniela Dutra, Nancy Philman, Dr. Michael Kane (University of Florida), Emily Massey, and Dr. Lawrence Zettler (Illinois College) was invaluable in the discovery of new records on the refuge. Great appreciation is also extended to Paul Martin Brown for his assistance in format and content of this flora. Financial support for this project was provided by the U.S. Fish and Wildlife Service to S.L.S. All photographs included in this flora were taken by the authors, unless otherwise indicated. The image of Harrisella porrecta was
103

Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

provided by Paul Martin Brown and the image of Sacoila lanceolata var. lanceolata forma albidaviridis was provided by Philip Kauth. References Ackerman, J.D. 1995. An orchid flora of Puerto Rico and the Virgin Islands. New York Botanical Garden, New York. Brown, P.M. and S.N. Folsom. 2005. Wild Orchids of Florida, updated and expanded edition. University Press of Florida, Gainesville. _____. 2002. Wild Orchids of Florida. University Press of Florida, Gainesville. Flora of North America Editorial Committee (eds.). 2002. Flora of North America north of Mexico, Orchidaceae. Pp. 490-651, vol. 26, New York & Oxford. Florida Panther National Wildlife Refuge (eds.). 2000. Florida Panther National Wildlife Refuge Comprehensive Conservation Plan. U.S. Fish and Wildlife Service, Florida. Hopper, S.D. 1997. An Australian perspective on plant conservation biology in practice. Pp. 255-78, in: Fiedler, P.L. & P.M. Kareiva (eds.) Conservation biology for the coming decade. Chapman Hall, New York. Johnson, T.R., S.L. Stewart, D. Dutra, M.E. Kane, and L. Richardson. 2007. Asymbiotic and symbiotic seed germination of Eulophia alta (Orchidaceae)—preliminary evidence for the symbiotic culture advantage. Plant Cell, Tissue and Organ Culture 90:313-23. Llamacho, J.A. and J.A. Larramendi. 2005. The Orchids of Cuba. Greta Editors, Spain. Luer, C.A. 1975. The Native Orchids of the United States and Canada, excluding Florida. New York Botanical Garden, New York. _____. 1972. The Native Orchids of Florida. New York Botanical Garden, New York. Menk, A. 2007. An extreme expression of color in Spiranthes odorata. North American Native Orchid Journal 13:26. Stewart, S.L. 2007. Integrated conservation of Florida Orchidaceae in the genera Habenaria and Spiranthes: model orchid conservation systems for the Americas. Ph.D. dissertation. University of Florida. Stewart, S.L. 2003. An orchid flora of the Florida Panther National Wildlife Refuge. U.S. Fish and Wildlife Service, Florida Panther National Wildlife Refuge, Florida. Stewart, S.L. and M.E. Kane. 2007. Orchid conservation in the Americas—lessons learned in Florida. Lankesteriana 7:382-87. Stewart, S.L., D. Dutra, P. Kauth, M.E. Kane, T. Johnson, N. Philman, and L. Richardson. 2007. Native Orchid Conservation on the Florida Panther National Wildlife Refuge, cooperative research funding projects on Eulophia alta, Cyrtopodium punctatum, Spiranthes odorata, and Bletia purpurea. U.S. Fish and Wildlife Service, Florida. Zettler, L.W., S.B. Poulter, K.I. McDonald, and S.L. Stewart. 2007. Conservation-driven propagation of an epiphytic orchid (Epidendrum nocturnum) with a mycorrhizal fungus. HortScience 42:135-39.

Scott L. Stewart, Ph.D.

PhytoTechnology Laboratories, LLC Lenexa, KS scott@phytotechlab.com

Larry W. Richardson

Florida Panther National Wildlife Refuge Naples, FL larry_richardson@fws.gov

104

SYNONYMS AND MISAPPLIED NAMES

SYNONYMS AND MISAPPLIED NAMES OF WILD ORCHIDS FOUND IN NORTH AMERICA NORTH OF MEXICO
The abundance of alternate names for many of our native North American orchids has often led to confusion among native orchid enthusiasts. Taxonomic opinion will always be with us, but understanding the place of synonyms and misapplied names helps to alleviate this confusion. The following reflects the current opinions of several generic specialists in the field. Synonyms for taxa at the forma rank are also given as many were originally described at the rank of variety and are better recognized as forma. Information has been taken, in part, from Brown and Folsom (2003, 2004, 2005, 2006a, 2006b, 2006c, 2007, 2008) and the International Plant Name Index (IPNI). Additional synonyms may be found in both of Luer‘s volumes on the orchids of the United States and Canada (1972, 1975) and, to a lesser extent, in Flora of North America North of Mexico volume 26, Orchidaceae (2002). The various publications, both in print and online, from Kartesz also contain additional synonyms. For a current listing see http://www.bonap.org. BASIONYM, a name originally applied to the species, is given first and most of the synonyms and misapplied names cited are from general usage in the 19th and 20th centuries; 18th century synonyms are given only when they have remained in the recent literature. SYNONYMS are alternate names used at various times by authors often embracing different generic concepts or a name that was published subsequent to the original publication. MISAPPLIED NAMES are incorrect names for a given species found in the continental United States or Canada; they are usually similar species often found in Mexico, Central America, South America, or the Caribbean. nom. nud. = without a name: used when a species is published or cited but without a Latin description or other critical part of the original publication rendering it invalid; technically it is not a synonym nom. illeg. = illegitimate name: used when a name is published that has previously been published for another taxon p. p. (pro parte) = in part; used when part of the name applies * non-native; ?=status questionable

Amerorchis rotundifolia (Banks) Hultén
BASIONYM: Orchis

Aplectrum hyemale (Mühlenberg in Willdenow) Nuttall
BASIONYM: Cymbidium hyemale Mühlenberg in Willdenow SYNONYM: Aplectrum spicatum Britton, Sterns &

rotundifolia Banks in Pursh forma angustifolia (Rousseau) P.M. Brown BASIONYM: Orchis rotundifolia (Banks in Pursh) forma angustifolia Rousseau forma beckettiae (Boivin) Hultén BASIONYM: Orchis rotundifolia (Banks in Pursh) forma beckettiae Boivin forma lineata (Mousley) Hultén BASIONYM: Orchis rotundifolia (Banks in Pursh) forma lineata Mousley Poggenberg

105

SYNONYMS AND MISAPPLIED NAMES forma pallidum (House) Barnhart BASIONYM: Aplectrum spicatum Britton, Sterns & Poggenberg forma pallidum House
BASIONYM: Carteria

Basiphyllaea corallicola (Small) Ames
corallicola Small
BASIONYM: Spiranthes

Beloglottis costaricensis (Reichenbach f.) Schlechter Bletia purpurea (Lamark) de Candolle
BASIONYM: Limodorum purpureum

costaricensis Reichenbach f.

Brassia caudata (Linnaeus) Lindley
BASIONYM: Epidendrum caudatum BASIONYM: Pleurothallis pachyrhachis

Lamark forma alba (Ariza-Julia & J. Jiménez Almonte) P.M. Brown BASIONYM: Bletia purpurea (Lamark) de Candolle var. alba Ariza-Julia & J. Jiménez Almonte Linnaeus

Bulbophyllum pachyrhachis (A. Richard) Grisebach Calopogon barbatus (Walter) Ames
A. Richard
BASIONYM: Ophrys barbata Walter SYNONYMS: Calopogon graminifolius Elliot in

Calopogon multiflorus Lindley
SYNONYMS:

Weatherby & Griscom Calopogon parviflorus Lindley Calopogon pulchellus (Salisbury) R. Brown var. graminifolius Elliot Limodorum parviflorum (Lindley) Nash Calopogon barbatus (Walter) Ames var. multiflorus (Lindley) Correll Limodorum multiflorum (Lindley) Mohr Limodorum pinetorum Small Limodorum pallidum (Chapman) Mohr

Calopogon pallidus Chapman
SYNONYM:

Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. tuberosus
BASIONYM: Cathea tuberosa (Linnaeus) Salisbury SYNONYMS: Calopogon pulchellus (Salisbury) R. Brown

Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. simpsonii (Small)
Magrath
BASIONYM: Limodorum simpsonii Small SYNONYM: Calopogon pulchellus (Salisbury) R. Brown BASIONYM: Calypso americana R. Brown in Aiton SYNONYM: Cytherea bulbosa (Linnaeus) House p.p.

Calopogon pulchellus (Salisbury) R. Brown var. latifolius (St. John) Fernald Limodorum pulchellum Salisbury Limodorum tuberosum Linnaeus

Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown) Luer Calypso bulbosa (Linnaeus) Oakes var. occidentalis (Holtzinger) Boivin
BASIONYM: Calypso bulbosa (Linnaeus) Oakes forma occidentalis Holzinger SYNONYMS: Calypso bulbosa Britton subsp. occidentalis (Holzinger) Calder &

var. simpsonii (Small) Ames

Cephalanthera austiniae (A. Gray) Heller
BASIONYM: Chloraea austinae A. Gray SYNONYMS: Cephalanthera oregana Reichenbach

Calypso occidentalis (Holzinger) A. Heller Cytherea bulbosa House var. occidentalis (Holzinger) Cockerell Cytherea occidentalis (Holzinger) A. Heller

Roy L. Taylor

Cleistes divaricata (Linnaeus) Ames

Eburophyton austinae (A. Gray) Heller Epipactis austinae (A. Gray) Wettstein Serapias austinae (A. Gray) A.A. Eaton

BASIONYM: Arethusa divaricata Linnaeus SYNONYM: Pogonia divaricata (Linnaeus) R. Brown

106

SYNONYMS AND MISAPPLIED NAMES

Cleistes bifaria (Fernald) Catling & Gregg

Coeloglossum viride (Linnaeus) Hartman var. viride
BASIONYM: Satyrium viride Linnaeus SYNONYMS: Coeloglossum viride (Linnaeus)

BASIONYM: Cleistes divaricata (Linnaeus) Ames var. bifaria Fernald SYNONYM: Pogonia bifaria (Fernald) P.M. Brown & Wunderlin

Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer
BASIONYM: Orchis bracteata Mühlenberg in Willdenow SYNONYMS: Coeloglossum bracteatum (Mühlenberg in Willdenow)

Hartman var. islandicum (Lindley) Schulze Dactylorhiza viridis (Linnaeus) R.M. Bateman, A. Pridgeon & M.W. Chase Habenaria viridis (Linnaeus) R. Brown in Aiton f.

Corallorhiza mertensiana Bongard
SYNONYMS:

Parlin Coeloglossum viride (Linnaeus) Hartman subsp. bracteatum (Mühlenberg in Willdenow) Hultén Habenaria bracteata (Mühlenberg in Willdenow) R. Brown in Aiton f. Habenaria viridis Linnaeus var. bracteata (Mühlenberg in Willdenow) Reichenbach in Gray Habenaria viridis Linnaeus var. interjecta Fernald Neottia mertensiana (Bongard) Kuntze Corallorhiza purpurea L.O. Williams Corallorhiza maculata (Rafinesque) Rafinesque subsp. mertensiana (Bongard) Calder & Taylor Corallorhiza vancouveriana Finet

Corallorhiza maculata (Rafinesque) Rafinesque var. maculata
BASIONYM: Cladorhiza maculata Rafinesque SYNONYM: Corallorhiza multiflora Nuttall

Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley) Ames
BASIONYM: Corallorhiza multiflora Nuttall var. occidentalis Lindley SYNONYM: Corallorhiza maculata (Rafinesque) Rafinesque subsp.

var. maculata forma flavida (Peck) Farwell BASIONYM: Corallorhiza multiflora Nuttall var. flava Peck

Corallorhiza maculata var. mexicana (Lindley) J.V. Freudenstein Corallorhiza odontorhiza (Willdenow) Poiret var. odontorhiza
BASIONYM: Corallorhiza

occidentalis (Lindley) Cockerell forma immaculata (Peck) Howell BASIONYM: Corallorhiza maculata (Rafinesque) Rafinesque var. immaculata Peck forma punicea (Bartholomew) Weatherby & Adams BASIONYM: Corallorhiza maculata (Rafinesque) Rafinesque var. punicea Bartholomew mexicana Lindley

Corallorhiza odontorhiza (Willdenow) Poiret var. pringlei (Greenman) Freudenstein Corallorhiza striata Lindley var. striata
SYNONYMS: BASIONYM: Corallorhiza

BASIONYM: Cymbidium odontorhizum Willdenow SYNONYM: Corallorhiza micrantha A. Chapman

pringlei Greenman

Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams
BASIONYM: Corallorhiza vreelandii Rydberg SYNONYMS: Corallorhiza bigelovii S. Watson

Corallorhiza macraei A. Gray Neottia striata (Lindley) Kuntze

Corallorhiza trifida Chatelain
SYNONYM:

Corallorhiza ochroleuca Rydberg forma flavida (Todsen & Todsen) P.M. Brown BASIONYM: Corallorhiza striata Lindley var. flavida T.A. Todsen & Todsen Corallorhiza corallorhiza (Linnaeus) Karsten forma verna (Nuttall) P.M. Brown BASIONYM: Corallorhiza verna Nuttall SYNONYM: Corallorhiza trifida Chatelain var. verna (Nuttall) Fernald
SYNONYMS:

Corallorhiza wisteriana Conrad

Corallorhiza odontorhiza sensu Chapman not Nuttall 107

SYNONYMS AND MISAPPLIED NAMES Corallorhiza unguiculata Rafinesque forma albolabia P.M. Brown SYNONYM: Corallorhiza wisteriana Conrad forma toleri S. Bentley
BASIONYM: Pelexia cranichoides Grisebach SYNONYMS: Beadlea cranichoides (Grisebach)

Cyclopogon cranichoides (Grisebach) Schlechter
Small Spiranthes cranichoides (Grisebach) Cogniaux

Cyclopogon elatus (Swartz) Schlechter
BASIONYM: Satryrium elatum Swartz SYNONYMS: Beadlea elata (Swartz) Small

Cypripedium acaule Aiton
SYNONYM: SYNONYM:

Spiranthes elata (Swartz) L.C. Richard Fissipes acaulis (Aiton) Small Criosanthes arietina (Aiton f.) House

Cypripedium arietinum R. Brown

Cypripedium kentuckiense C.F. Reed
Cypripedium daultonii V. Soukup nom. nud.
SYNONYM: SYNONYM:

Cypripedium fasciculatum Kellogg in S. Watson Cypripedium montanum Douglas in Lindley Cypripedium parviflorum Salisbury var. parviflorum
SYNONYM: Cypripedium

Cypripedium knightae A. Nelson Cypripedium occidentale Watson

Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight
BASIONYM: Cypripedium pubescens Willdenow SYNONYMS: Cypripedium calceolus Linnaeus var. BASIONYM: Cypripedium pubescens Willdenow var. makasin Farwell SYNONYM: Cypripedium calceolus Linnaeus var. parviflorum (Salisbury) Fernald

calceolus Linnaeus var. parviflorum (Salisbury) Fernald p.p.

p.p.

Cypripedium passerinum Richardson forma minganense (Victorin) P.M. Brown Cypripedium reginae Walter
SYNONYM: Cypripedium SYNONYMS: BASIONYM: Cypripedium passerinum

planipetalum (Fernald) Victorin & Rousseau Cypripedium calceolus Linnaeus var. pubescens (Willdenow) Correll Cypripedium flavescens de Candolle Cypripedium parviflorum var. planipetalum Fernald Cypripedium pubescens Willdenow Cypripedium veganum Cockerell & Barber MISAPPLIED NAME: Cypripedium calceolus Linnaeus Richardson var. minganense Victorin

Cypripedium yatabeanum Makino

spectabile Salisbury

Cyrtopodium macrobulbon (La Llave & Lexarza) G.A. Romero & Carnevali Cyrtopodium polyphyllum (Vellozo) Pabst in F. Barrios Cyrtopodium punctatum (Linnaeus) Lindley
BASIONYM: Epidendrum punctatum BASIONYM: Orchis BASIONYM: Epidendrum polyphyllum Vellozo MISAPPLIED NAME: Cyrtopodium andersonii (Lambert in BASIONYM: Epidendrum macrobulbon

Cypripedium guttatum subsp. yatabeanum (Makino) Hultén Cypripedium guttatum var. yatabeanum (Makino) Pfitzer La Llave & Lexarza

Andrews) R. Brown in Aiton

Dactylorhiza aristata (Fischer in Lindley) Soó Dactylorhiza aristata (Fischer in Lindley) Soó var. kodiakensis Luer forma perbracteata
(Lepage) P.M. Brown BASIONYM: Orchis aristata forma perbracteata Lepage aristata Fischer in Lindley

Linnaeus

108

SYNONYMS AND MISAPPLIED NAMES
SYNONYM: Dactylorhiza

Dactylorhiza majalis (Reichenbach f.) Summerhayes (Druce) D.M. Moore & Soó subsp. praetermissa var. junialis (Vermeulen) Senghas *?
BASIONYM: Orchis latifolius Linnaeus var. junialis Vermeulen SYNONYMS: Dactylorhiza junialis Vermeulen

aristata (Fischer ex Lindley) Soó forma perbracteata (Lepage) Catling

Dactylorhiza majalis (Reichenbach f.) Summerhayes var. praetermissa (Druce) D.M.
Moore & Soó *?
BASIONYM: Orchis praetermissa Druce SYNONYM: Dactylorhiza praetermissa (Druce) Soó

Dactylorhiza majalis (Reichenbach f.) P.E Hunt & Summerhayes Dactylorhiza praetermissa (Druce) Soó var. junialis (Vermeulen) Senghas MISAPPLIED NAMES: Dactylorhiza cf. fuchsii (Druce) Soó Dactylorhiza comosa subsp. majalis (Reichenbach f.) P.D. Sell Dactylorhiza maculata (Linnaeus) Soó Orchis purpurella T. & T.A. Stephenson

Deiregyne confusa Garay
SYNONYMS:

var. praetermissa forma albiflora (Druce) P.M. Brown BASIONYM: Orchis praetermissa Druce var. albiflora Druce Spiranthes confusa (Garay) Kartesz & Gandhi Funkiella confusa (Garay) Szlachetko Schiedeella confusa (Garay) Espejo & López-Ferrari MISAPPLIED NAMES: Deiregyne durangensis (Ames & Schweinfurth) Garay Spiranthes durangensis Ames & Schweinfurth
BASIONYM: Aeranthes porrectus Reichenbach f. SYNONYMS: Campylocentrum porrectum Reichenbach f.

Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten
Harrisella porrecta (Reichenbach f.) Fawcett & Rendle

Dendrophylax lindenii (Lindley) Bentham in Rolfe
BASIONYM: Angraecum lindenii Lindley SYNONYMS: Polyradicion lindenii (Lindley) Garay

Dichromanthus cinnabarinus (La Llave & Lexarza) Garay
BASIONYM: Neottia cinnabarina La Llave & Lexarza SYNONYMS: Spiranthes cinnabarina (La Llave & Lexarza) Hemsley

Polyrrhiza lindenii (Lindley) Cogniaux

Dichromanthus michuacanus (La Llave & Lexarza) Salazar & Soto-Arenas
BASIONYM: Neottia michuacana La Llave & Lexarza SYNONYMS: Spiranthes michuacana (La Llave & Lexarza)

Stenorrhynchos cinnabarina (La Llave & Lexarza) Lindley

Eltroplectris calcarata (Swartz) Garay & Sweet
BASIONYM: Neottia calcarata Swartz SYNONYMS: Centrogenium setaceum (Lindley) Schlechter

Hemsley Stenorrhynchos michuacanum (La Llave & Lexarza) Lindley

Encyclia rufa (Lindley) Britton & Millspaugh
BASIONYM: Epidendrum rufum Lindley SYNONYM: Encyclia bahamensis (Grisebach) Britton & MISAPPLIED NAME: Epidendrum bahamense Grisebach BASIONYM: Epidendrum tampense

Pelexia setacea Lindley

Millspaugh

Encyclia tampensis (Lindley) Small Epidendrum acuñae Dressler
Lindley

Epidendrum amphistomum A. Richard
MISAPPLIED NAME: Epidendrum anceps

MISAPPLIED NAME: Epidendrum blancheanum

Urban

Epidendrum magnoliae Mühlenberg var. magnoliae
109

Jacquin

SYNONYMS AND MISAPPLIED NAMES
SYNONYM:

Epidendrum magnoliae Mühlenberg var. mexicanum (L.O. Williams) P.M. Brown Epidendrum floridense Hágsater
BASIONYM: Epidendrum conopseum

Epidendrum conopseum R. Brown

R. Brown var. mexicanum L.O. Williams

MISAPPLIED NAMES: Epidendrum difforme

Epipactis atrorubens (Hoffman) Besser* Epipactis gigantea Douglas in Hooker
SYNONYMS: BASIONYM: Serapias

Jacquin Epidendrum umbellatum Swartz Neolehmannia difformis (Jacquin) Pabst

latifolia Hudson (rank unspecified) atrorubens Hoffman

Epipactis helleborine (Linnaeus) Cranz*

Amesia gigantea (Douglas) A. Nelson & Macbride Helleborine gigantea (Douglas) Druce Peramium giganteum (Douglas) Salisbury

BASIONYM: Serapias helleborine Linnaeus SYNONYMS: Amesia latifolia (Linnaeus) A. Nelson

Eulophia alta (Linnaeus) Fawcett & Rendle
BASIONYM: Limodorum altum

& J.F. MacBryde Epipactis latifolia (Linnaeus) Allioni forma alba (Webster) Boivin BASIONYM: Epipactis latifolia Allioni forma alba Webster forma monotropoides (Mousley) Scoggin BASIONYM: Amesia latifolia A. Nelson & J.F. MacBryde forma monotropoides Mousley forma variegata (Webster) Boivin BASIONYM: Epipactis latifolia (Linnaeus) Allioni forma variegata Webster Linnaeus

Galeandra bicarinata G.A. Romero & P.M. Brown Galearis spectabilis (Linnaeus) Rafinesque
BASIONYM: Orchis spectabilis Linnaeus SYNONYM: Galeorchis spectabilis (Linnaeus) MISAPPLIED NAME: Galeandra

beyrichii Reichenbach f.

Goodyera oblongifolia Rafinesque
SYNONYMS:

Rydberg forma gordinierii (House) Whiting & Catling BASIONYM: Galeorchis spectabilis (Linnaeus) Rydberg forma gordinierii House SYNONYM: Galearis spectabilis (Linnaeus) Rafinesque forma albiflora (Ulke) C.F. Reed Orchis spectabilis Linnaeus forma albiflora Ulke forma lilacina (Ames) P.M. Brown BASIONYM: Orchis spectabilis Linnaeus forma lilacina Ames SYNONYM: Galearis spectabilis (Linnaeus) Rafinesque forma willeyi (Seymour) P.M. Brown Goodyera decipiens (Hooker) E.T. Hubbard Peramium decipiens (Hooker) Piper Spiranthes decipiens Hooker forma reticulata (Boivin) P.M. Brown BASIONYM: Goodyera oblongifolia Rafinesque var. reticulata Boivin

Goodyera pubescens (Willdenow) R. Brown Goodyera repens (Linnaeus) R. Brown
BASIONYM: Satyrium repens Linnaeus SYNONYM: Goodyera pubescens (Willdenow)

BASIONYM: Neottia pubescens Willdenow SYNONYM: Peramium pubescens (Willdenow) MacMillan

R. Brown var. repens (R. Brown) Alphonse Wood ophioides Fernald

forma ophioides (Fernald) P.M. Brown

Goodyera tesselata Loddiges
SYNONYM:

BASIONYM: Goodyera repens (Linnaeus) R. Brown var. SYNONYM: Peramium ophioides (Fernald) Rydberg

Peramium tesselatum (Loddiges) A. Heller 110

SYNONYMS AND MISAPPLIED NAMES

Govenia floridana P.M. Brown
MISAPPLIED NAME: Govenia BASIONYM: Orchis

Gymnadenia conopsea (Linnaeus) R. Brown* Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata
BASIONYM: Orchis clavellata Michaux SYNONYMS: Habenaria clavellata (Michaux) Sprengel

utriculata (Swartz) Lindley

conopsea Linnaeus

Gymnadeniopsis clavellata (Michaux) Rydberg var. ophioglossoides (Fernald) W.J.
Schrenk
BASIONYM: Habenaria clavellata (Michaux) Sprengel var. ophioglossoides Fernald SYNONYM: Platanthera clavellata (Michaux) Luer var. ophioglossoides (Fernald) P.M. Brown BASIONYM: Orchis integra Nuttall SYNONYMS: Habenaria integra (Nuttall) Sprengel

Platanthera clavellata (Michaux) Luer forma slaughteri (P.M. Brown) P.M. Brown BASIONYM: Platanthera clavellata (Michaux) Luer var. clavellata forma slaughteri P.M. Brown forma wrightii (Olive) P.M. Brown BASIONYM: Habenaria clavellata (Michaux) Sprengel var. wrightii Olive

Gymnadeniopsis integra (Nuttall) Rydberg
Platanthera integra (Nuttall) Luer

Gymnadeniopsis nivea (Nuttall) Rydberg
BASIONYM: Orchis nivea Nuttall SYNONYMS: Habenaria nivea (Nuttall) Sprengel

Habenaria macroceratitis Willdenow
SYNONYM: SYNONYMS: Habenaria

Platanthera nivea (Nuttall) Luer

Habenaria odontopetala Reichenbach f.

Habenaria quinqueseta var. macroceratitis (Willdenow) Luer

Habenaria quinqueseta (Michaux) Eaton
BASIONYM: Orchis quinqueseta Michaux SYNONYMS: Habenaria michauxii Nuttall

garberi Porter Habenaria strictissima Reichenbach f. var. odontopetala (Reichenbach f.) L.O. Williams Habenella odontopetala (Reichenbach f.) Small MISAPPLIED NAME: Habenaria floribunda Lindley

Habenaria repens Nuttall
SYNONYMS:

Habenaria simpsonii Small Platanthera michauxii (Nuttall) Wood Habenaria nuttallii Small Platanthera repens (Nuttall) Wood

Harrisella porrecta (Reichenbach f.) Fawcett & Rendle
BASIONYM: Campylocentrum porrectum Reichenbach f. SYNONYM: Dendrophylax porrectus (Reichenbach f.) Carlsward & MISAPPLIED NAME: Harrisella filiformis (Swartz) Cogniaux BASIONYM: Corallorhiza grandiflora A. Richard SYNONYM: Hexalectris mexicana Greenman

Whitten

Hexalectris grandiflora (A. Richard & Galeotti) L.O. Williams
& Galeotti

Hexalectris spicata (Walter) Barnhardt

Hexalectris spicata var. arizonica (S. Watson) Catling & Engel Ionopsis utricularioides (Swartz) Lindley Isotria medeoloides (Pursh) Rafinesque
111
BASIONYM: Epidendrum utricularioides BASIONYM: Corallorhiza

BASIONYM: Arethusa spicata Walter SYNONYM: Hexalectris aphylla (Nuttall) Rafinesque

arizonica S. Watson

Swartz

SYNONYMS AND MISAPPLIED NAMES
BASIONYM: Arethusa medeoloides Pursh SYNONYMS: Isotria affinis (Austin in A. Gray) Rydberg

Isotria verticillata (Mühlenberg in Willdenow) Rafinesque Lepanthopsis melanantha (Reichenbach f.) Ames Liparis elata Lindley
SYNONYM: BASIONYM: Pleurothallis melanantha Reichenbach f. SYNONYM: Lepanthes harrissii Fawcett & Rendell MISAPPLIED NAME: Liparis nervosa BASIONYM: Arethusa verticillata Mühlenberg in Willdenow SYNONYM: Pogonia verticillata (Mühlenberg in Willdenow) Nuttall

Pogonia affinis Austin in A. Gray

Listera auriculata Wiegand

(Thunberg) Lindley

Listera australis Lindley
SYNONYM: SYNONYMS:

Neottia auriculata (Wiegand) Szlachetko forma trifolia (Lepage) Lepage BASIONYM: Listera borealis Morong forma trifolia Lepage Neottia australis (Lindley) Szlachetko

Listera banksiana Lindley

Listera borealis Morong
SYNONYM: BASIONYM: Ophrys SYNONYM: Neottia BASIONYM: Listera SYNONYMS:

Listera caurina Piper Listera retusa Suksdorf Neottia caurina (Piper) Szlachetko Neottia borealis (Morong) Szlachetko cordata Linnaeus cordata (Linnaeus) Richard

Listera cordata (Linnaeus) R. Brown var. cordata Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hultén
nephrophylla Rydberg Listera cordata (Linnaeus) R. Brown subsp. nephrophylla (Rydberg) A. & D. Löve Neottia nephrophylla (Rydberg) Szlachetko Bluff & Fingerhut

Listera ovata (Linnaeus) R. Brown* Listera smallii Wiegand
SYNONYMS: BASIONYM: Ophrys ovata Linnaeus SYNONYM: Neottia ovata (Linnaeus)

Malaxis abieticola Salazar & Soto Arenas Malaxis bayardii Fernald
MISAPPLIED NAME: Malaxis tenuis (S.

Listera reniformis Small Neottia smallii (Wiegand) Szlachetko

Watson) Ames

Malaxis brachypoda (Gray) Fernald

Malaxis unifolia var. bayardii nom. nud.

BASIONYM: Microstylis brachypoda A. Gray SYNONYM: Malaxis monophyllos (Linnaeus) Swartz var. brachypoda MISAPPLIED NAME: Microstylis monophyllos (Linnaeus) Lindley

(A. Gray) Morris & Eames

Malaxis corymbosa (S. Watson) Kuntze Malaxis diphyllos Chamisso
BASIONYM: Microstylis corymbosa S. Watson SYNONYM: Achroanthes corymbosa Greene

forma bifolia (Mousley) Fernald BASIONYM: Malaxis monophyllos (Linnaeus) Swartz forma bifolia Mousley

SYNONYM: Malaxis monophyllos (Linnaeus) Swartz var. diphyllos MISAPPLIED NAMES: Malaxis monophyllos (Linnaeus) Swartz

(Chamisso) Luer

Malaxis paludosa Swartz

Microstylis monophyllos (Linnaeus) Lindley

112

SYNONYMS AND MISAPPLIED NAMES
SYNONYM:

Malaxis porphyrea (Ridley) Kuntze
BASIONYM: Microstylis porphyrea Ridley MISAPPLIED NAMES: Malaxis ehrenbergii

Hammarbya paludosa Kuntze

Malaxis spicata Swartz
SYNONYMS:

Malaxis wendtii Salazar

(Reichenbach f.) Kuntze

Malaxis soulei L.O. Williams
SYNONYMS:

Achroanthes floridana (Chapman) Greene Malaxis floridana (Chapman) O. Kuntze Microstylis floridana Chapman Microstylis spicata (Swartz) Lindley Achroanthes montana Greene Malaxis macrostachys (Lexarza) Kuntze Malaxis montana (Englemann) Kuntze Microstylis macrostachys Lexarza Microstylis montana Englemann Achroanthes monophylla (Lindley) Greene Microstylis unifolia (Michaux) Britton, Sterns & Poggenberg Tamayorkis wendtii (Salazar) R. González & Szlachetko Leon

Malaxis unifolia Michaux
SYNONYMS:

Malaxis wendtii Salazar
SYNONYM:

Maxillaria parviflora (Poeppig & Endlicher) Garay Mesadenus lucayanus (Britton) Schlechter
BASIONYM: Ibidium lucayanum Britton MISAPPLIED NAME: Mesadenus polyanthus BASIONYM: Scaphyglottis parviflora Poeppig & Endlicher SYNONYM: Maxillaria conferta (Grisebach) Schweinfurth in

Microthelys rubrocallosa (Robinson & Greenman) Garay
BASIONYM: Spiranthes rubrocallosa Robinson & Greenman SYNONYMS: Galeottiella rubrocallosa (Robinson & Greenman)

(Reichenbach f.) Schlechter

Oeceoclades maculata (Lindley) Lindley* Pelexia adnata (Swartz) Sprengel
BASIONYM: Satyrium adnatum Swartz SYNONYM: Spiranthes adnata (Swartz) BASIONYM: Limodorum tankervilleae BASIONYM: Angraecum

Szlachetko Schiedeella rubrocallosa (Robinson & Greenman) Burns-Balogh maculatum Lindley

Phaius tankervilleae (Aiton) Blume
Aiton

Bentham in Fawcett

Piperia candida R. Morgan & J. Ackerman Piperia colemanii Morgan & Glicenstein Piperia cooperi (S. Watson) Rydberg
BASIONYM: Habenaria cooperi S. Watson SYNONYM: Platanthera cooperi (S. Watson) BASIONYM: Platanthera SYNONYMS: Habenaria SYNONYM:

Platanthera candida (R. Morgan & J. Ackerman) R.M. Bateman nom. illeg. Platanthera colemanii (Morgan & Glicenstein) R.M. Bateman R.M. Bateman

Piperia elegans (Lindley) Rydberg subsp. elegans

elegans Lindley elegans (Lindley) Bolander Habenaria elegans (Lindley) Bolander var. maritima (Greene) Ames Habenaria greenei Jepson Habenaria maritima Greene Habenaria unalascensis (Sprengel) S. Watson var. maritima (Greene) Correll Piperia maritima (Greene) Rydberg Piperia multiflora Rydberg 113

SYNONYMS AND MISAPPLIED NAMES Platanthera unalascensis (Sprengel) Kurtz subsp. maritima (Greene) de Filipps

Piperia elongata Rydberg
SYNONYMS:

Piperia leptopetala Rydberg
SYNONYM:

Habenaria unalascensis (Sprengel) S. Watson var. elata (Jepson) Correll Piperia elegans var. elata (Jepson) Luer Piperia lancifolia Rydberg Piperia longispica Durand Platanthera elongata (Rydberg) R.M. Bateman Platanthera unalascensis (Sprengel) Kurtz subsp. elata (Jepson) Taylor & MacBryde Platanthera leptopetala (Rydberg) R.M. Bateman

Piperia michaelii (Greene) Rydberg

BASIONYM: Habenaria michaelii Greene SYNONYMS: Piperia elongata Rydberg subsp.

Piperia transversa Suksdorf
SYNONYM:

michaelii (Greene) Ackerman Platanthera michaelii (E. Greene) R.M. Bateman Platanthera transversa (Suksdorf) R.M. Bateman

Piperia unalascensis (Sprengel) Rydberg
BASIONYM: Spiranthes unalascensis Sprengel SYNONYMS: Habenaria unalascensis (Sprengel)

Piperia yadonii R. Morgan & J. Ackerman
SYNONYMS:

S. Watson Platanthera unalascensis (Sprengel) Kurtz MISAPPLIED NAME: Platanthera foetida Geyer in Hooker Platanthera yadonii (R. Morgan & J. Ackerman) R.M. Bateman Habenaria californica Grinnell nom. nud.
MISAPPLIED NAMES: Habenaria

Platanthera aquilonis Sheviak

Platanthera blephariglottis (Willdenow) Lindley
BASIONYM: Orchis blephariglottis Willdenow SYNONYMS: Blephariglottis blephariglottis (Willdenow)

hyperborea (Linnaeus) R. Brown in Aiton Platanthera hyperborea (Linnaeus) Lindley forma alba (Light) P.M. Brown BASIONYM: Platanthera hyperborea (Linnaeus) Lindley forma alba M.H.S. Light Rydberg

Platanthera brevifolia (Greene) Kranzlein
BASIONYM: Habenaria

Habenaria blephariglottis (Willdenow) Hooker brevifolia Greene

Platanthera chapmanii (Small) Luer emend. Folsom
BASIONYM: Blephariglottis chapmanii Small SYNONYMS: Habenaria xchapmanii (Small) Platanthera xchapmanii (Small) Luer

Ames

Platanthera chorisiana (Chamisso) Reichenbach f.
BASIONYM: Habenaria chorisiana Chamisso SYNONYMS: Limnorchis chorisiana (Chamisso)

Platanthera ciliaris (Linnaeus) Lindley
Habenaria ciliaris (Linnaeus) R. Brown

J.P. Anderson Pseudodiphryllum chorisianum (Chamisso) Nevski

BASIONYM: Orchis ciliaris Linnaeus SYNONYMS: Blephariglottis ciliaris (Linnaeus) Rydberg

Platanthera conspicua (Nash) P.M. Brown
BASIONYM: Habenaria conspicua Nash SYNONYMS: Blephariglottis conspicua (Nash)

Platanthera convallariifolia Fischer in Lindley
SYNONYM:

Small Habenaria blephariglottis var. conspicua (Nash) Ames Platanthera blephariglottis (Willdenow) Lindley var. conspicua (Nash) Luer Limnorchis convallariaefolius (Fischer) Rydberg 114

SYNONYMS AND MISAPPLIED NAMES
MISAPPLIED NAME: Habenaria

Platanthera cristata (Michaux) Lindley
Habenaria cristata (Michaux) R. Brown

borealis Chamisso var. viridiflora Chamisso

BASIONYM: Orchis cristata Michaux SYNONYMS: Blephariglottis cristata (Michaux) Rafinesque

Platanthera dilatata (Pursh) Lindley
BASIONYM: Orchis dilatata Pursh SYNONYMS: Limnorchis dilatata (Pursh)

Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour
BASIONYM: Habenaria borealis Chamisso var. albiflora Chamisso SYNONYMS: Habenaria dilatata (Pursh) Hooker var. albiflora (Chamisso)

Rydberg in Britton Piperia dilatata (Pursh) Szlachetko & P. Rutkowski Platanthera dilatata (Pursh) Lindley var. angustifolia Hooker

Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer
BASIONYM: Platanthera SYNONYMS: Habenaria

Correll Piperia dilatata (Pursh) Szlachetko & P. Rutkowski var. albiflora (Chamisso) Szlachetko & P. Rutkowski leucostachys Lindley dilatata (Pursh) Hooker var. leucostachys (Lindley) Ames Habenaria leucostachys (Lindley) S. Watson Limnorchis leucostachys (Lindley) Rydberg R. Brown in Sprengel

Platanthera flava (Linnaeus) Lindley var. flava
BASIONYM: Orchis flava Linnaeus SYNONYMS: Habenaria flava (Linnaeus)

Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer
BASIONYM: Habenaria herbiola R. Brown in Aiton SYNONYMS: Habenaria flava (Linnaeus) R. Brown in

Platanthera grandiflora (Bigelow) Lindley

Sprengel var. herbiola (R. Brown) Ames & Correll Habenaria flava (Linnaeus) R. Brown in Sprengel var. virescens sensu Fernald forma lutea (Boivin) Whiting & Catling BASIONYM: Habenaria flava (Linnaeus) R. Brown forma lutea Louis-Marie in Boivin

BASIONYM: Orchis grandiflora Bigelow SYNONYMS: Blephariglottis grandiflora (Bigelow) Rydberg

Platanthera hookeri (Torrey) Lindley
BASIONYM: Habenaria hookeri Torrey SYNONYMS: Habenaria hookeriana Torrey

Fimbriella psycodes (Linnaeus) Butzin var. grandiflora (Bigelow) Butzin Habenaria fimbriata (Dryander) R. Brown in Aiton Habenaria grandiflora (Bigelow) Torrey Habenaria psycodes (Linnaeus) Sprengel var. grandiflora (Bigelow) A. Gray forma albiflora (Rand & Redfield) Catling BASIONYM: Habenaria fimbriata (Dryander) R. Brown in Aiton var. albiflora Rand & Redfield forma mentotonsa (Fernald) P.M. Brown BASIONYM: Habenaria fimbriata (Dryander) R. Brown in Aiton forma mentotonsa Fernald in A. Gray Lysias hookeriana Rydberg in Britton Orchis hookeriana Oakes forma abbreviata (Fernald) P.M. Brown BASIONYM: Habenaria hookeri Torrey var. abbreviata Fernald SYNONYMS: Platanthera hookeri (Torrey ex A. Gray) Lindley var. abbreviata (Fernald) Catling Platanthera hookeri (Torrey in A. Gray) Lindley var. abbreviata (Fernald) W.J. Schrenk forma oblongifolia (J.A. Paine) P.M. Brown BASIONYM: Platanthera hookeri (Torrey) Lindley var. oblongifolia J.A. Paine
BASIONYM: Orchis huronensis Nuttall SYNONYMS: Habenaria hyperborea (Linnaeus) Habenaria xmedia (Rydberg) Niles

Platanthera huronensis (Nuttall) Lindley

R. Brown var. huronensis (Nuttall) Farwell

Limnorchis media Rydberg

115

SYNONYMS AND MISAPPLIED NAMES Platanthera dilatata (Pursh) Lindley var. chlorantha Hultén Platanthera hyperborea (Linnaeus) Lindley var. huronensis (Nuttall) Luer Platanthera xmedia (Rydberg) Luer R. Brown in Aiton Limnorchis hyperborea (Linnaeus) Rydberg Butzin Habenaria lacera (Michaux) R. Brown

Platanthera hyperborea (Linnaeus) Lindley
BASIONYM: Orchis hyperborea Linnaeus SYNONYMS: Habenaria hyperborea (Linnaeus)

Platanthera lacera (Michaux) G. Don
BASIONYM: Orchis lacera Michaux SYNONYM: Fimbriella lacera (Michaux) F.

Platanthera leucophaea (Nuttall) Lindley
BASIONYM: Orchis leucophaea Nuttall SYNONYMS: Fimbriella leucophaea (Nuttall) Farwell

Platanthera limosa Lindley
SYNONYMS:

Habenaria leucophaea (Nuttall) A. Gray

Platanthera macrophylla (Goldie) P.M. Brown
BASIONYM: Habenaria macrophylla Goldie SYNONYMS: Habenaria orbiculata (Pursh) Hooker

Habenaria limosa (Lindley) Hemsley Habenaria thurberi A. Gray Limnorchis arizonica Rydberg

Platanthera obtusata (Banks in Pursh) Lindley
BASIONYM: Orchis obtusata Banks in Pursh SYNONYMS: Habenaria obtusata (Banks in Pursh)

forma macrophylla (Goldie) F. Morris & E.A. Eames Habenaria orbiculata (Pursh) Hooker var. macrophylla (Goldie) B. Boivin Platanthera orbiculata (Pursh) Lindley var. macrophylla (Goldie) Luer Lysias macrophylla (Goldie) House

Platanthera orbiculata (Pursh) Lindley
BASIONYM: Orchis orbiculata Pursh SYNONYMS: Habenaria orbiculata (Pursh)

Richards Lysiella obtusata Rydberg forma collectanea (Fernald) P.M. Brown BASIONYM: Habenaria obtusata (Banks in Pursh) Richards var. collectanea Fernald Torrey Habenaria orbiculata (Pursh) Torrey var. menziesii (Lindley) Fernald Lysias orbiculata Rydberg Lysias menziesii (Lindley) Rydberg Platanthera menziesii Lindley forma lehorsii (Fernald) P.M. Brown BASIONYM: Habenaria orbiculata (Pursh) Torrey var. lehorsii Fernald forma longifolia (Clute) P.M. Brown BASIONYM: Habenaria orbiculata (Pursh) Torrey var. longifolia Clute forma pauciflora (Jennings) P.M. Brown BASIONYM: Lysias orbiculata Rydberg var. pauciflora Jennings forma trifolia (Mousley) P.M. Brown BASIONYM: Habenaria orbiculata (Pursh) Torrey var. trifolia Mousley
BASIONYM: Habenaria peramoena A. Gray SYNONYM: Blephariglottis peramoena (Gray) Rydberg

Platanthera peramoena (A. Gray) A. Gray
Fimbriella peramoena (A. Gray) F. Butzin

Platanthera praeclara Sheviak & Bowles
SYNONYMS:

Platanthera psycodes (Linnaeus) Lindley
BASIONYM: Orchis

Fimbriella praeclara (Sheviak & M.L. Bowles) Szlachetko & Rutkowski Habenaria leucophaea (Nuttall) A. Gray var. praeclara (Sheviak & Bowles) Cronquist psycodes Linnaeus

116

SYNONYMS AND MISAPPLIED NAMES
SYNONYMS:

Platanthera purpurascens (Rydberg) Sheviak & W.F. Jennings
BASIONYM: Limnorchis purpurascens Rydberg SYNONYM: Habenaria purpurascens (Rydberg) Tidestrom

Blephariglottis psycodes (Linnaeus) Rydberg Fimbriella psycodes (Linnaeus) Butzin Habenaria psycodes (Linnaeus) Sprengel forma albiflora (R. Hoffman) Whiting & Catling BASIONYM: Habenaria psycodes (Linnaeus) Sprengel forma albiflora Hoffman forma ecalcarata (Bryan) P.M. Brown BASIONYM: Habenaria psycodes (Linnaeus) Sprengel var. ecalcarata Bryan forma fernaldii (Rousseau & Rouleau) P.M. Brown BASIONYM: Habenaria psycodes (Linnaeus) Sprengel var. fernaldii Rousseau & Rouleau forma varians (Bryan) P.M. Brown BASIONYM: Habenaria psycodes (Linnaeus) Sprengel var. varians Bryan

Platanthera sparsiflora (S. Watson) Schlechter
BASIONYM: Habenaria sparsiflora S. Watson SYNONYMS: Limnorchis laxiflora Rydberg

Habenaria hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Ames Platanthera hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Luer

Platanthera stricta Lindley
SYNONYMS:

Limnorchis sparsiflora (S. Watson) Rydberg Habenaria saccata Greene Limnorchis stricta (Lindley) Rydberg Platanthera gracilis Lindley Platanthera hyperborea (Linnaeus) Lindley var. viridiflora (Chamisso) Kitamura Platanthera hyperborea (Linnaeus) Lindley var. viridiflora (Chamisso) Luer Platanthera saccata (Greene) Hultén Limnorchis ensifolia Rydberg Platanthera sparsiflora (S. Watson) Schlechter var. ensifolia (Rydberg) Luer

Platanthera tescamnis Sheviak & W.F. Jennings
SYNONYMS:

Platanthera tipuloides (Linnaeus) Lindley var. behringiana (Rydberg) Hultén Platanthera xandrewsii (Niles) Luer
BASIONYM: Limnorchis behringiana Rydberg SYNONYM: Habenaria behringiana (Rydberg) Ames

Fimbriella andrewsii (White) Butzin Fimbriella lacera (Michaux) Butzin var. terrae-novae (Fernald) Butzin Platanthera andrewsii (White) Luer Platanthera lacera var. terrae-novae (Fernald) Luer Platanthera xbeckneri P.M. Brown MISAPPLIED NAME: Platanthera xcanbyi (Ames) Luer Platanthera xbicolor (Rafinesque) Luer SYNONYM: Habenaria xbicolor (Rafinesque) Beckner Platanthera xcanbyi (Ames) Luer BASIONYM: Habenaria xcanbyi Ames SYNONYM: Blephariglottis canbyi (Ames) W. Stone Platanthera xlueri P.M. Brown MISAPPLIED NAME: Platanthera xbicolor (Rafinesque) Luer XPlatanthopsis vossii (Case) P.M. Brown BASIONYM: Platanthera xvossii Case

BASIONYM: Habenaria xandrewsii M. White in G.G. Niles SYNONYMS:: Blephariglottis xandrewsii (White) House

Platythelys latifolia (Linnaeus) Garay & Ormerod
BASIONYM: Satytium latifolium Linnaeus SYNONYMS: Erythrodes sagreana (A. Richard)

Physurus sagreanus A. Richard

Leon 117

SYNONYMS AND MISAPPLIED NAMES Platythelys sagreana (A. Richard) Garay

Platythelys querceticola (Lindley) Garay Pleurothallis gelida Lindley
SYNONYM: BASIONYM: Arethusa BASIONYM: Physurus querceticola Lindley SYNONYM: Erythrodes querceticola (Lindley)

Ames

Pogonia ophioglossoides (Linnaeus) Ker-Gawler

Stelis gelida (Lindley) Pridgeon & M.W. Chase

Polystachya concreta (Jacquin) Garay & Sweet Ponthieva brittoniae Ames
SYNONYM: BASIONYM: Arethusa BASIONYM: Epidendrum concretum Jacquin MISAPPLIED NAME: Polystachya flavescens (Lindley)

ophioglossoides Linnaeus forma brachypogon (Fernald) P.M. Brown BASIONYM: Pogonia ophioglossoides (Linnaeus) Ker-Gawler var. brachypogon Fernald J. K. Small

Ponthieva racemosa (Walter) Mohr
racemosa Walter
BASIONYM: Cranichis oligantha

Ponthieva racemosa (Walter) C. Mohr var. brittonae (Ames) Luer

Prescottia oligantha (Swartz) Lindley Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins
BASIONYM: Epidendrum cochleatum Linnaeus var. triandrum Ames SYNONYMS: Anacheilium cochleatum (Linnaeus) Small var. triandrum

Swartz

Prosthechea pygmaea (Hooker) W.E. Higgins
BASIONYM: Epidendrum pygmaceum Hooker SYNONYMS: Encyclia pygmaea (Hooker) Dressler

(Ames) Sauleda et al. Encyclia cochleata (Linnaeus) Dressler subsp. triandra (Ames) Hágsater forma albidoflava (P.M. Brown) P.M. Brown SYNONYM: Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler forma albidoflava P.M. Brown

Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E.
Higgins
BASIONYM: Epidendrum erythronioides Small SYNONYMS: Encyclia bidentata (Lindley) Hágsater

Hormidium pygmaeum (Hooker) Bentham & Hooker f.

Pseudorchis straminea (Fernald) Soó
BASIONYM: Habenaria straminea Fernald SYNONYMS: Habenaria albida (Linnaeus)

& Soto Arenas subsp. erythronioides (Small) Hágsater Encyclia boothiana (Lindley) Dressler var. erythronioides (Small) Luer Epicladium boothianum (Lindley) Small var. erythronioides (Small) Acuña Pseudencyclia boothiana (Lindley) V.P. Castro & Chiron R. Brown var. straminea (Fernald) Morris & Eames Leucorchis albida subsp. straminea (Fernald) A. Löve Pseudorchis albida (Linnaeus) Löve & Löve subsp. straminea (Fernald) Löve & Löve

Pteroglossaspis ecristata (Fernald) Rolfe
BASIONYM: Cyrtopodium ecristatum Fernald SYNONYM: Eulophia ecristata (Fernald) Ames

Sacoila lanceolata (Aublet) Garay var. lanceolata
BASIONYM: Limodorum lanceolatum Aublet SYNONYMS: Spiranthes lanceolata (Aublet) Leon

Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Sauleda, Wunderlin & Hansen
BASIONYM: Spiranthes

Stenorrhynchos lanceolatum (Aublet) Richard in Sprengel Stenorrhynchos orchoides (Swartz) L.C. Richard var. lanceolata (Aublet) Garay forma albidaviridis Catling & Sheviak MISAPPLIED NAME: Spiranthes lanceolata (Aublet) Garay var. luteoalba (Reichenbach f.) Luer lanceolata (Aublet) Leon var. paludicola Luer 118

SYNONYMS AND MISAPPLIED NAMES
SYNONYM: Stenorrhynchos cinnabarinms (La Llave BASIONYM: Neottia squamulosa Kunth SYNONYMS: Sacoila lanceolata (Aublet)

Sacoila squamulosa (Kunth) Garay

& Lexarza) Lindley var. paludicola (Luer) W. J. Schrenk

Schiedeella arizonica P.M. Brown
MISAPPLIED NAMES: Schiedeella

Garay var. squamulosa (Kunth) Szlachetko Spiranthes squamulosa (Kunth) Leon Stenorrhynchos squamulosum (Kunth) Sprengel fauci-sanguinea (Dod) Burns-Balogh Schiedeella parasitica (A. Richard & Galeotti) Schlechter Spiranthes fauci-sanguinea Dod Spiranthes parasitica A. Richard & Galeotti Spiranthes gracilis (Bigelow) Beck var. brevilabris (Lindley) Correll Ames

Spiranthes brevilabris Lindley
SYNONYM:

Spiranthes casei var. casei Catling & Cruise Spiranthes cernua (Linnaeus) L.C. Richard
BASIONYM: Ophrys cernua Linnaeus SYNONYMS: Ibidium cernuum (Linnaeus) House MISAPPLIED NAME: Spiranthes intermedia

Spiranthes delitescens Sheviak Spiranthes diluvialis Sheviak
SYNONYM:

Triorchis cernua (Linnaeus) Nieuwland

MISAPPLIED NAME: Spiranthes graminea

Lindley

Spiranthes floridana (Wherry) Cory emend. P.M. Brown
BASIONYM: Ibidium floridanum Wherry SYNONYMS: Spiranthes brevilabris Lindley

Spiranthes romanzoffiana Chamisso var. diluvialis (Sheviak) S.L. Welsh

Spiranthes lacera Rafinesque var. lacera
BASIONYM: Neottia

var. floridana (Wherry) Luer Spiranthes gracilis var. floridana (Wherry) Correll lacera Rafinesque

Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer
BASIONYM: Neottia gracilis Bigelow SYNONYMS: Ibidium beckii (Lindley)

Spiranthes laciniata (Small) Ames
Triorchis laciniata House

House Ibidium gracile (Bigelow) House Spiranthes beckii Lindley Spiranthes gracilis (Bigelow) Beck

BASIONYM: Gyrostachys laciniata Small SYNONYM: Ibidium laciniatum (Small) House

Spiranthes longilabris Lindley
SYNONYMS:

Spiranthes lucida (H.H. Eaton) Ames
Spiranthes plantaginea Rafinesque Triorchis plantaginea Nieuwland

Ibidium longilabre (Lindley) House Spiranthes brevifolia Chapman Triorchis longilabris House House

BASIONYM: Neottia lucida H.H. Eaton SYNONYMS: Ibidium plantagineum (Rafinesque)

Spiranthes ochroleuca (Rydberg) Rydberg

BASIONYM: Gyrostachys ochroleuca Rydberg in Britton SYNONYMS: Spiranthes cernua (Linnaeus) L.C. Richard Spiranthes xsteigeri Correll

var. ochroleuca (Rydberg) Ames

Triorchis cernua Nieuwland var. ochroleuca Farwell Triorchis ochroleuca Nieuwland 119

SYNONYMS AND MISAPPLIED NAMES

Spiranthes odorata (Nuttall) Lindley

BASIONYM: Neottia odorata Nuttall SYNONYMS: Ibidium odoratum (Nuttall) House

Spiranthes ovalis Lindley
SYNONYM: SYNONYMS:

Spiranthes cernua (Linnaeus) L.C. Richard var. odorata (Nuttall) Correll Triorchis odorata (Nuttall) Nieuwland Ibidium ovale (Lindley) House

Spiranthes porrifolia Lindley

Spiranthes praecox (Walter) S. Watson
BASIONYM: Limodorum praecox Walter SYNONYM: Ibidium praecox (Walter) House

Gyrostachys porrifolia (Lindley) Kuntze Ibidium porrifolium Rydberg Spiranthes romanzoffiana Chamisso var. porrifolia (Lindley) Ames & Correll

Spiranthes romanzoffiana Chamisso
SYNONYMS:

Triorchis praecox (Walter) Nieuwland

Spiranthes torta (Thunberg) Garay & Sweet
BASIONYM: Ophrys torta Thunberg SYNONYM: Ibidium tortile (Swartz) House MISAPPLIED NAME: Spiranthes tortilis (Swartz) SYNONYMS:

Gyrostachys romanzoffiana (Chamisso) MacMillan Gyrostachys striata Rydberg Ibidium strictum (Rydberg) House Spiranthes stricta (Rydberg) A. Nelson in J.M. Coulter & A. Nelson Triorchis romanzoffiana (Chamisso) Nieuwland Triorchis stricta (Rydberg) Nieuwland

Spiranthes tuberosa Rafinesque

Richard

Spiranthes vernalis Engelmann & A. Gray
SYNONYMS:

Ibidium beckii House Spiranthes grayi Ames Spiranthes tuberosa var. grayi (Ames) Fernald Triorchis grayi (Ames) Nieuwland

Ibidium vernale (Engelmann & A. Gray) House Gyrostachys vernalis Kuntze Triorchis vernalis House Spiranthes xmeridionalis P.M. Brown Spiranthes xaustralis P.M. Brown nom..illeg.

Tipularia discolor (Pursh) Nuttall
BASIONYM: Orchis discolor Pursh SYNONYM: Tipularia unifolia Britton,

Tolumnia bahamense (Nash in Britton & Millspaugh) G. J. Braem Trichocentrum carthagenense (Jacquin) M.W. Chase & N.H. Williams Trichocentrum luridum (Lindley) M.W. Chase & N.H. Williams
BASIONYM: Oncidium SYNONYM: BASIONYM: Epidendrum carthagenense Jacquin SYNONYM: Lophiaris carthagenensis (Jacquin) Braem BASIONYM: Oncidium bahamense Nash in Britton & Millspaugh MISAPPLIED NAME: Tolumnia variegata (Swartz) G. J. Braem

Sterns & Poggenberg

Trichocentrum undulatum (Swartz) Ackerman & Chase
BASIONYM: Epidendrum undulatum Swartz SYNONYMS: Oncidium undulatum (Swartz) Salisbury

luridum Lindley Lophiaris lurida (Lindley) Braem

Trichocentrum maculatum (Aublet) M. Chase & N.A. Williams nom. illeg. 120

SYNONYMS AND MISAPPLIED NAMES forma flavovirens (P.M. Brown) P.M. Brown BASIONYM: Oncidium undulatum (Swartz) Salisbury forma flavovirens P.M. Brown SYNONYM: Trichocentrum maculatum (Aublet) M.W. Chase & N.H. Williams forma flavovirens (P.M. Brown) P.M. Brown
SYNONYM:

Triphora amazonica Schlechter
Triphora latifolia G. Luer

Triphora gentianoides (Swartz) Ames & Schlechter*
BASIONYM: Limodorum gentianoides Swartz SYNONYMS: Pogonia cubensis Reichenbach f.

Triphora rickettii Luer

Pogonia gentianoides (Swartz) Sprengel Triphora cubensis (Reichenbach f.) Ames yucatanensis Ames

Triphora trianthophoros (Swartz) Rydberg
BASIONYM: Arethusa trianthophoros Swartz SYNONYMS: Arethusa pendula Mühlenberg

MISAPPLIED NAME: Triphora

Tropidia polystachya (Swartz) Ames Vanilla barbellata Reichenbach f. Vanilla dilloniana Correll Vanilla inodora Schiede
SYNONYMS: MISAPPLIED NAME: SYNONYM: BASIONYM: Serapias polystachya Swartz SYNONYM: Tropidia eatonii Ames

in Willdenow Pogonia pendula (Mühlenberg in Willdenow) Lindley Pogonia trianthophoros (Swartz) Britton, Sterns & Poggenberg Triphora pendula (Mühlenberg in Willdenow) Nuttall

Vanilla articulata Northrop

Vanilla eggersii Rolfe

Vanilla planifolia Andrews
SYNONYM:

Epidendrum vanilla Linnaeus p.p. Vanilla mexicana Miller

Vanilla fragrans (Salisbury) Ames Vanilla vanilla (Linnaeus) Britton

CROSS-REFERENCES FOR SYNONYMS AND MISAPPLIED NAMES = synonym and/or basionym ≠ misapplied name Achroanthes corymbosa Greene = Malaxis corymbosa (S. Watson) Kuntze Achroanthes floridana (Chapman) Greene = Malaxis spicata Swartz Achroanthes monophylla (Lindley) Greene = Malaxis unifolia Lindley Achroanthes montana Greene = Malaxis soulei L.O. Williams Aeranthes porrectus Reichenbach f. = Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten Amesia gigantea (Douglas) A. Nelson & Macbride = Epipactis gigantea Douglas in Hooker Amesia latifolia (Linnaeus) A. Nelson & J.F. MacBryde = Epipactis helleborine (Linnaeus) Cranz Amesia latifolia A. Nelson & J.F. MacBryde forma monotropoides Mousley = Epipactis helleborine (Linnaeus) Cranz forma monotropoides (Mousley) Scoggin Anacheilium cochleatum (Linnaeus) Small var. triandrum (Ames) Sauleda et al. = Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins Angraecum lindenii Lindley = Dendrophylax lindenii (Lindley) Bentham in Rolfe Angraecum maculatum Lindley = Oeceoclades maculata (Lindley) Lindley Aplectrum spicatum Britton, Sterns & Poggenberg = Aplectrum hyemale (Mühlenberg in Willdenow) Nuttall
121

SYNONYMS AND MISAPPLIED NAMES

Arethusa divaricata Linnaeus = Cleistes divaricata (Linnaeus) Ames Arethusa medeoloides Pursh = Isotria medeoloides (Pursh) Rafinesque Arethusa ophioglossoides Linnaeus = Pogonia ophioglossoides (Linnaeus) Ker-Gawler Arethusa pendula Mühlenberg in Willdenow = Triphora trianthophoros (Swartz) Rydberg Arethusa racemosa Walter = Ponthieva racemosa (Walter) Mohr Arethusa trianthophoros Swartz = Triphora trianthophoros (Swartz) Rydberg Arethusa verticillata Mühlenberg in Willdenow = Isotria verticillata (Mühlenberg in Willdenow) Rafinesque Beadlea cranichoides (Grisebach) Small = Cyclopogon cranichoides (Grisebach) Schlechter Beadlea elata (Swartz) Small = Cyclopogon elatus (Swartz) Schlechter Blephariglottis blephariglottis (Willdenow) Rydberg = Platanthera blephariglottis (Willdenow) Lindley Blephariglottis canbyi (Ames) W. Stone = Platanthera xcanbyi (Ames) Luer Blephariglottis chapmanii Small = Platanthera chapmanii (Small) Luer emend. Folsom Blephariglottis ciliaris (Linnaeus) Rydberg = Platanthera ciliaris (Linnaeus) Lindley Blephariglottis conspicua (Nash) Small = Platanthera conspicua (Nash) P.M. Brown Blephariglottis cristata (Michaux) Rafinesque = Platanthera cristata (Michaux) Lindley Blephariglottis flaviflora Rafinesque = Platanthera ciliaris (Linnaeus) Lindley Blephariglottis grandiflora (Bigelow) Rydberg = Platanthera grandiflora (Bigelow) Lindley Blephariglottis peramoena (Gray) Rydberg = Platanthera peramoena (A. Gray) A. Gray Blephariglottis psycodes (Linnaeus) Rydberg = Platanthera psycodes (Linnaeus) Lindley Bletia purpurea (Lamark) de Candolle var. alba Ariza-Julia & J. Jiménez Almonte = Bletia purpurea (Lamark) de Candolle forma alba (Ariza-Julia & J. Jiménez Almonte) P.M. Brown Calopogon barbatus (Walter) Ames var. multiflorus (Lindley) Correll = Calopogon multiflorus Lindley Calopogon graminifolius Elliot in Weatherby & Griscom = Calopogon barbatus (Walter) Ames Calopogon parviflorus Lindley = Calopogon barbatus (Walter) Ames Calopogon pulchellus (Salisbury) R. Brown = Calopogon tuberosus (Linnaeus) Britton, Sterns & Calopogon pulchellus (Salisbury) R. Brown var. graminifolius Elliot = Calopogon barbatus (Walter) Ames Calopogon pulchellus (Salisbury) R. Brown var. latifolius (St. John) Fernald = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg Calopogon pulchellus (Salisbury) R. Brown var. simpsonii (Small) Ames = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. simpsonii (Small) Magrath Calypso americana R.Brown in Aiton = Calypso bulbosa Britton var. americana (R. Brown) Luer Calypso bulbosa subsp. occidentalis (Holtzman) Calder & Taylor = Calypso bulbosa (Linnaeus) Oakes var. occidentalis (Holtzman) Boivin Campylocentrum porrectum Reichenbach f. = Harrisella porrecta (Reichenbach f.) Fawcett & Rendle Carteria corallicola Small = Basiphyllaea corallicola (Small) Ames Cathea tuberosa (Linnaeus) Salisbury = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg Centrogenium setaceum (Lindley) Schlechter = Eltroplectris calcarata (Swartz) Garay & Sweet Cephalanthera oregana Reichenbach = Cephalanthera austiniae (A. Gray) Heller Chloraea austinae Gray = Cephalanthera austiniae (A. Gray) Heller Cladorhiza maculata Rafinesque = Corallorhiza maculata (Rafinesque) Rafinesque var. maculata Cleistes divaricata (Linnaeus) Ames var. bifaria Fernald = Cleistes bifaria (Fernald) Catling & Gregg Coeloglossum bracteatum (Mühlenberg in Willdenow) Parlin = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer Coeloglossum viride (Linnaeus) Hartman subsp. bracteatum (Mühlenberg in Willdenow) Hultén = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer Coeloglossum viride (Linnaeus) Hartman var. islandicum (Lindley) Schulze = Coeloglossum viride (Linnaeus) Hartman var. viride
122

SYNONYMS AND MISAPPLIED NAMES

Corallorhiza arizonica S. Watson = Hexalectris spicata var. arizonica (S. Watson) Catling & Engel Corallorhiza bigelovii S. Watson = Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams Corallorhiza corallorhiza (Linnaeus) Karsten = Corallorhiza trifida Chatelain Corallorhiza grandiflora Richard & Galeotti = Hexalectris grandiflora (Richard & Galeotti) L.O. Williams Corallorhiza macraei A. Gray = Corallorhiza striata Lindley Corallorhiza maculata subsp. mertensiana (Bongard) Calder & Taylor = Corallorhiza mertensiana Bongard Corallorhiza maculata subsp. occidentalis (Lindley) Cockerell = Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley) Ames Corallorhiza mexicana Lindley = Corallorhiza maculata var. mexicana (Lindley) J.V. Freudenstein Corallorhiza micrantha A. Chapman = Corallorhiza odontorhiza (Willdenow) Nuttall Corallorhiza multiflora Nuttall = Corallorhiza maculata (Rafinesque) Rafinesque var. maculata Corallorhiza multiflora Nuttall var. occidentalis Lindley = Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley) Ames Corallorhiza odontorhiza sensu Chapman not Nuttall = Corallorhiza wisteriana Conrad Corallorhiza pringlei Greenman = Corallorhiza odontorhiza (Willdenow) Poiret var. pringlei (Greenman) Freudenstein Corallorhiza purpurea L.O. Williams = Corallorhiza mertensiana Bongard Corallorhiza striata Lindley var. flavida T.A. Todsen & Todsen = Corallorhiza striata Lindley forma flavida (Todsen & Todsen) P.M. Brown Corallorhiza trifida Chatelain var. verna (Nuttall) Fernald = Corallorhiza trifida Chatelain forma verna (Nuttall) P.M. Brown Corallorhiza unguiculata Rafinesque = Corallorhiza wisteriana Conrad Corallorhiza vancouveriana Finet = Corallorhiza mertensiana Bongard Corallorhiza vreelandii Rydberg = Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams Corallorhiza wisteriana Conrad forma toleri S. Bentley = Corallorhiza wisteriana Conrad forma albolabia P.M. Brown Cranichis oligantha Swartz = Prescottia oligantha (Swartz) Lindley Criosanthes arietina (Aiton f.) House = Cypripedium arietinum R. Brown Cymbidium hyemale Mühlenberg in Willdenow = Aplectrum hyemale (Mühlenberg in Willdenow) Nuttall Cymbidium odontorhizum Willdenow = Corallorhiza odontorhiza (Willdenow) Poiret var. odontorhiza Cypripedium calceolus Linnaeus ≠ Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium calceolus Linnaeus var. parviflorum (Salisbury) Fernald = Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak Cypripedium calceolus Linnaeus var. parviflorum (Salisbury) Fernald p.p. = Cypripedium parviflorum Salisbury var. parviflorum Cypripedium calceolus Linnaeus var. planipetalum (Fernald) Victorin & Rousseau = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium calceolus Linnaeus var. pubescens (Willdenow) Correll = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium daultonii V. Soukup nom. nud. = Cypripedium kentuckiense C.F. Reed Cypripedium flavescens de Candolle = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium guttatum subsp. yatabeanum (Makino) Hultén = Cypripedium yatabeanum Makino Cypripedium guttatum var. yatabeanum (Makino) Pfitzer = Cypripedium yatabeanum Makino Cypripedium knightae A. Nelson = Cypripedium fasciculatum Kellogg in S. Watson
123

SYNONYMS AND MISAPPLIED NAMES

Cypripedium occidentale Watson = Cypripedium montanum Douglas in Lindley Cypripedium parviflorum var. planipetalum Fernald = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium passerinum Richardson var. minganense Victorin = Cypripedium passerinum Richardson forma minganense (Victorin) P.M. Brown Cypripedium pubescens var. makasin Farwell = Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak Cypripedium pubescens Willdenow = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cypripedium spectabile Salisbury = Cypripedium reginae Walter Cypripedium veganum Cockerell & Barber = Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight Cyrtopodium andersonii (Lambert in Andrews) R. Brown in W.T. Aiton ≠Cyrtopodium polyphyllum (Vellozo) Pabst in F. Barrios Cyrtopodium ecristatum Fernald = Pteroglossaspis ecristata (Fernald) Rolfe Cytherea bulbosa (Linnaeus) House = Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown) Luer Dactylorhiza aristata (Fischer ex Lindley) Soó forma perbracteata (Lepage) Catling = Dactylorhiza aristata (Fischer ex Lindley) Soó var. kodiakensis Luer forma perbracteata (Lepage) P.M. Brown Dactylorhiza cf. fuchsii (Druce) Soó ≠ Dactylorhiza majalis subsp. praetermissa (Druce) D.M. Moore & Soó subsp var. junialis (Vermeulen) Senghas Dactylorhiza comosa subsp. majalis (Reichenbach f.) P.D. Sell ≠ Dactylorhiza majalis subsp. praetermissa (Druce) D.M. Moore & Soó var. junialis (Vermeulen) Senghas Dactylorhiza maculata (Linnaeus) Soó ≠ Dactylorhiza majalis subsp. praetermissa var. junialis (Vermeulen) Senghas Dactylorhiza majalis (Reichenbach f.) P.E Hunt & Summerhayes = Dactylorhiza majalis subsp. praetermissa (Druce) D.M. Moore & Soó var. junialis (Vermeulen) Senghas Dactylorhiza praetermissa (Druce) Soó var. junialis (Vermeulen) Senghas = Dactylorhiza majalis subsp. praetermissa (Druce) D.M. Moore & Soó var. junialis (Vermeulen) Senghas Dactylorhiza praetermissa (Druce) Soó var. praetermissa = Dactylorhiza majalis (Reichenbach) Summerhayes var. praetermissa (Druce) D.M. Moore & Soó Dactylorhiza viridis (Linnaeus) R.M. Bateman, A. Pridgeon & M.W. Chase = Coeloglossum viride (Linnaeus) Hartman Deiregyne durangensis (Ames & Schweinfurth) Garay ≠ Deiregyne confusa Garay Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten = Harrisella porrecta (Reichenbach f.) Fawcett & Rendle Eburophyton austinae (A. Gray) Heller = Cephalanthera austiniae (A. Gray) Heller Encyclia bahamensis (Grisebach) Britton & Millspaugh = Encyclia rufa (Lindley) Britton & Millspaugh Encyclia bidentata (Lindley) Hágsater & Soto Arenas subsp. erythronioides (Small) Hágsater = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Encyclia boothiana (Lindley) Dressler var. erythronioides (Small) Luer = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Encyclia cochleata (Linnaeus) Dressler subsp. triandra (Ames) Hágsater = Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler = Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins
124

SYNONYMS AND MISAPPLIED NAMES

Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler forma albidoflava P.M. Brown = Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins forma albidoflava (P.M. Brown) P.M. Brown Encyclia pygmaea (Hooker) Dressler = Prosthechea pygmaea (Hooker) W.E. Higgins Epicladium boothianum (Lindley ) Small var. erythronioides (Small) Acuña = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Epidendrum anceps Jacquin ≠ Epidendrum amphistomum A. Richard Epidendrum bahamense Grisebach ≠ Encyclia rufa (Lindley) Britton & Millspaugh Epidendrum blancheanum Urban ≠ Epidendrum acuñae Dressler Epidendrum boothianum Lindley = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Epidendrum carthagenense Jacquin = Trichocentrum carthagenense (Jacquin) M.W. Chase & N.H. Williams Epidendrum caudatum Linnaeus = Brassia caudata (Linnaeus) Lindley Epidendrum cochleatum Linnaeus var. triandrum Ames = Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins Epidendrum concretum Jacquin = Polystachya concreta (Jacquin) Garay & Sweet Epidendrum conopseum R. Brown = Epidendrum magnoliae Mühlenberg Epidendrum conopseum R. Brown var. mexicanum L.O. Williams = Epidendrum magnoliae Mühlenberg var. mexicanum (L.O. Williams) P.M. Brown Epidendrum difforme Jacquin ≠ Epidendrum floridense Hágsater Epidendrum erythronioides Small = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Epidendrum macrobulbon La Llave & Lexarza = Cyrtopodium macrobulbon (La Llave & Lexarza) G.A. Romero & Carnevali Epidendrum polyphyllum Vellozo = Cyrtopodium polyphyllum (Vellozo) Pabst in F. Barrios Epidendrum punctatum Linnaeus = Cyrtopodium punctatum (Linnaeus) Lindley Epidendrum pygmaceum Hooker = Prosthechea pygmaea (Hooker) W.E. Higgins Epidendrum rufum Lindley = Encyclia rufa (Lindley) Britton & Millspaugh Epidendrum tampense Lindley = Encyclia tampensis (Lindley) Small Epidendrum umbellatum Swartz ≠ Epidendrum floridense Hágsater Epidendrum undulatum Swartz = Trichocentrum undulatum (Swartz) Ackerman & Chase Epidendrum utricularioides Swartz = Ionopsis utricularioides (Swartz) Lindley Epipactis austinae (A. Gray) Wettstein = Cephalanthera austiniae (A. Gray) Heller Epipactis latifolia (Linnaeus) Allioni = Epipactis helleborine (Linnaeus) Cranz Epipactis latifolia Allioni forma alba Webster = Epipactis helleborine (Linnaeus) Cranz forma alba (Webster) Boivin Epipactis latifolia (Linnaeus) Allioni forma variegata Webster = Epipactis helleborine (Linnaeus) Cranz forma variegata (Webster) Boivin Erythrodes querceticola (Lindley) Ames = Platythelys querceticola (Lindley) Garay Erythrodes sagreana (A. Richard) Leon = Platythelys latifolia (Linnaeus ) Garay & Ormerod Eulophia ecristata (Fernald) Ames = Pteroglossaspis ecristata (Fernald) Rolfe Fimbriella andrewsii (White in Niles) Butzin = Platanthera xandrewsii (White in Niles) Luer Fimbriella lacera (Michaux) Butzin var. terrae-novae (Fernald) Butzin = Platanthera xandrewsii (White in Niles) Luer Fimbriella lacera (Michaux) F. Butzin = Platanthera lacera (Michaux) G. Don Fimbriella leucophaea (Nuttall) Farwell = Platanthera leucophaea (Nuttall) Lindley
125

SYNONYMS AND MISAPPLIED NAMES

Fimbriella peramoena (A. Gray) F. Butzin = Platanthera peramoena (A. Gray) A. Gray Fimbriella praeclara (Sheviak & M.L. Bowles) Szlachetko & Rutkowski = Platanthera praeclara Sheviak & Bowles Fimbriella psycodes (Linnaeus) Butzin = Platanthera psycodes (Linnaeus) Lindley Fimbriella psycodes (Linnaeus) Butzin var. grandiflora (Bigelow) Butzin = Platanthera grandiflora (Bigelow) Lindley Fissipes acaulis (Aiton) Small = Cypripedium acaule Aiton Funkiella confusa (Garay) Szlachetko = Deiregyne confusa Garay Galearis spectabilis (Linnaeus) Rafinesque forma albiflora (Ulke) C.F. Reed = Galearis spectabilis (Linnaeus) Rafinesque forma gordinierii (House) Whiting & Catling Galearis spectabilis (Linnaeus) Rafinesque forma willeyi (Seymour) P.M. Brown = Galearis spectabilis (Linnaeus) Rafinesque forma lilacina (Ames) P.M. Brown Galeandra beyrichii Reichenbach f. ≠ Galeandra bicarinata G.A. Romero & P.M. Brown Galeorchis spectabilis (Linnaeus) Rydberg = Galearis spectabilis (Linnaeus) Rafinesque Galeorchis spectabilis (Linnaeus) Rydberg forma gordinierii House = Galearis spectabilis (Linnaeus) Rafinesque forma gordinierii (House) Catling & Whiting Galeottiella rubrocallosa (Robinson & Greenman) Szlachetko = Microthelys rubrocallosa (Robinson & Greenman) Garay Goodyera decipiens (Hooker) E.T. Hubbard = Goodyera oblongifolia Rafinesque Goodyera oblongifolia Rafinesque var. reticulata Boivin = Goodyera oblongifolia Rafinesque forma reticulata (Boivin) P.M. Brown Goodyera pubescens (Willdenow) R. Brown var. repens (R. Brown) Alphonse Wood = Goodyera repens (Linnaeus) R. Brown Goodyera repens (Linnaeus) R. Brown var. ophioides Fernald = Goodyera repens (Linnaeus) R. Brown forma ophioides (Fernald) P.M. Brown Govenia utriculata (Swartz) Lindley ≠ Govenia floridana P.M. Brown Gymnadenia albida subsp. straminea (Fernald) B. Ljtnant = Pseudorchis straminea (Fernald) Soó Gyrostachys laciniata Small = Spiranthes laciniata (Small) Ames Gyrostachys ochroleuca Rydberg in Britton = Spiranthes ochroleuca (Rydberg) Rydberg Gyrostachys porrifolia (Lindley) Kuntze = Spiranthes porrifolia Lindley Gyrostachys romanzoffiana (Chamisso) MacMillan = Spiranthes romanzoffiana Chamisso Gyrostachys striata Rydberg = Spiranthes romanzoffiana Chamisso Gyrostachys vernalis Kuntze = Spiranthes vernalis Engelmann & A. Gray Habenaria albida (Linnaeus) R. Brown var. straminea (Fernald) Morris & Eames = Pseudorchis straminea (Fernald) Soó Habenaria behringiana (Rydberg) Ames = Platanthera tipuloides (Linnaeus) Lindley var. behringiana (Rydberg) Hultén Habenaria blephariglottis (Willdenow) Hooker = Platanthera blephariglottis (Willdenow) Lindley Habenaria blephariglottis (Willdenow) Hooker var. integrilabia Correll = Platanthera integrilabia (Correll) Luer Habenaria blephariglottis var. conspicua (Nash) Ames = Platanthera conspicua (Nash) P.M. Brown Habenaria borealis Chamisso var. albiflora Chamisso = Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour Habenaria borealis Chamisso var. viridiflora Chamisso = Platanthera stricta Lindley Habenaria borealis Chamisso var. viridiflora Chamisso ≠ Platanthera convallariifolia Fischer in Lindley Habenaria bracteata (Mühlenberg in Willdenow) R. Brown in Aiton f. = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer
126

SYNONYMS AND MISAPPLIED NAMES

Habenaria brevifolia Greene = Platanthera brevifolia (Greene) Kranzlein Habenaria californica Grinnell nom. nud. = Piperia yadonii R. Morgan & J. Ackerman Habenaria chorisiana Chamisso = Platanthera chorisiana (Chamisso) Reichenbach f. Habenaria ciliaris (Linnaeus) R. Brown = Platanthera ciliaris (Linnaeus) Lindley Habenaria clavellata (Michaux) Sprengel = Gymnadeniopsis clavellata (Michaux) Rydberg Habenaria clavellata (Michaux) Sprengel var. wrightii Olive = Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata forma wrightii (Olive) P.M. Brown Habenaria clavellata (Michaux) Sprengel var. ophioglossoides Fernald = Gymnadeniopsis clavellata (Michaux) Rydberg var. ophioglossoides (Fernald) W.J. Schrenk Habenaria conspicua Nash = Platanthera conspicua (Nash) P.M. Brown Habenaria cooperi S. Watson = Piperia cooperi (S. Watson) Rydberg Habenaria correlliana Cronquist = Platanthera integrilabia (Correll) Luer Habenaria cristata (Michaux) R. Brown = Platanthera cristata (Michaux) Lindley Habenaria dilatata (Pursh) Hooker var. albiflora (Chamisso) Correll = Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour Habenaria dilatata (Pursh) Hooker var. leucostachys (Lindley) Ames = Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer Habenaria elegans (Lindley) Bolander = Piperia elegans (Lindley) Rydberg subsp. elegans Habenaria elegans (Lindley) Bolander var. maritima (Greene) Ames = Piperia elegans (Lindley) Rydberg subsp. elegans Habenaria fimbriata (Dryander) R. Brown in Aiton = Platanthera grandiflora (Bigelow) Lindley Habenaria flava (Linnaeus) R. Brown in Sprengel = Platanthera flava (Linnaeus) Lindley var. flava Habenaria flava (Linnaeus) R. Brown forma lutea Louis-Marie in Boivin = Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer forma lutea (Boivin) Whiting & Catling Habenaria flava (Linnaeus) R. Brown in Sprengel var. herbiola (R. Brown in Aiton) Ames & Correll = Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer Habenaria flava (Linnaeus) R. Brown in Sprengel var. virescens sensu Fernald = Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer Habenaria floribunda Lindley ≠ Habenaria odontopetala Reichenbach f. Habenaria garberi Porter = Habenaria odontopetala Reichenbach f. Habenaria grandiflora (Bigelow) Torrey = Platanthera grandiflora (Bigelow) Lindley Habenaria greenei Jepson = Piperia elegans (Lindley) Rydberg subsp. elegans Habenaria herbiola R. Brown in Aiton = Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer Habenaria hookeri Torrey = Platanthera hookeri (Torrey) Lindley Habenaria hookeri Torrey var. abbreviata Fernald = Platanthera hookeri (Torrey) Lindley forma abbreviata (Fernald) P.M. Brown Habenaria hookeriana Torrey in A. Gray = Platanthera hookeri (Torrey) Lindley Habenaria hyperborea (Linnaeus) R. Brown in Aiton = Platanthera hyperborea (Linnaeus) Lindley Habenaria hyperborea (Linnaeus) R. Brown in Aiton ≠ Platanthera aquilonis Sheviak Habenaria hyperborea (Linnaeus) R. Brown var. huronensis (Nuttall) Farwell = Platanthera huronensis (Nuttall) Lindley Habenaria hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Ames = Platanthera purpurascens (Rydberg) Sheviak & W. Jennings Habenaria integra (Nuttall) Sprengel = Gymnadeniopsis integra (Nuttall) Rydberg Habenaria lacera (Michaux) R. Brown = Platanthera lacera (Michaux) G. Don Habenaria lacera (Michaux) R. Brown var. terrae-novae Fernald = Platanthera xandrewsii (Niles) Luer
127

SYNONYMS AND MISAPPLIED NAMES

Habenaria leucophaea (Nuttall) A. Gray = Platanthera leucophaea (Nuttall) Lindley Habenaria leucophaea (Nuttall) A. Gray var. praeclara (Sheviak & Bowles) Cronquist = Platanthera praeclara Sheviak & Bowles Habenaria leucostachys (Lindley) S. Watson = Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer Habenaria limosa Hemsley = Platanthera limosa Lindley Habenaria macrophylla Goldie = Platanthera macrophylla (Goldie) P.M. Brown Habenaria maritima Greene = Piperia elegans (Lindley) Rydberg subsp. elegans Habenaria michaelii Greene = Piperia michaelii (Greene) Rydberg Habenaria michauxii Nuttall = Habenaria quinqueseta (Michaux) Eaton Habenaria nivea (Nuttall) Sprengel = Gymnadeniopsis nivea (Nuttall) Rydberg Habenaria nuttallii Small = Habenaria repens Nuttall Habenaria obtusata (Banks in Pursh) Richards = Platanthera obtusata (Banks in Pursh) Lindley Habenaria obtusata (Banks in Pursh) Richards var. collectanea Fernald = Platanthera obtusata (Banks in Pursh) Lindley forma collectanea (Fernald) P.M. Brown Habenaria orbiculata (Pursh) Torrey = Platanthera orbiculata (Pursh) Lindley Habenaria orbiculata (Pursh) Hooker forma macrophylla (Goldie) F. Morris & E.A. Eames = Platanthera macrophylla (Goldie) P.M. Brown Habenaria orbiculata (Pursh) Hooker var. macrophylla (Goldie) B. Boivin = Platanthera macrophylla (Goldie) P.M. Brown Habenaria orbiculata (Pursh) Torrey var. lehorsii Fernald = Platanthera orbiculata (Pursh) Lindley forma lehorsii (Fernald) P.M. Brown Habenaria orbiculata (Pursh) Torrey var. longifolia Clute = Platanthera orbiculata (Pursh) Lindley forma longifolia (Clute) P.M. Brown Habenaria orbiculata (Pursh) Torrey var. menziesii (Lindley) Fernald= Platanthera orbiculata (Pursh) Lindley Habenaria orbiculata (Pursh) Torrey var. trifolia Mousley = Platanthera orbiculata (Pursh) Lindley forma trifolia (Mousley) P.M. Brown Habenaria peramoena A. Gray = Platanthera peramoena (A. Gray) A. Gray Habenaria psycodes (Linnaeus) Sprengel = Platanthera psycodes (Linnaeus) Lindley Habenaria psycodes (Linnaeus) Sprengel forma albiflora Hoffman = Platanthera psycodes (Linnaeus) Lindley forma albiflora (R. Hoffman) Whiting & Catling Habenaria psycodes (Linnaeus) Sprengel var. ecalcarata Bryan = Platanthera psycodes (Linnaeus) Lindley forma ecalcarata (Bryan) P.M. Brown Habenaria psycodes (Linnaeus) Sprengel var. fernaldii Rousseau & Rouleau = Platanthera psycodes (Linnaeus) Lindley forma fernaldii (Rousseau & Rouleau) P.M. Brown Habenaria psycodes (Linnaeus) Sprengel var. varians Bryan = Platanthera psycodes (Linnaeus) Lindley forma varians (Bryan) P.M. Brown Habenaria psycodes (Linnaeus) Sprengel var. grandiflora (Bigelow) A. Gray = Platanthera grandiflora (Bigelow) Lindley Habenaria purpurascens (Rydberg) Tidestrom = Platanthera purpurascens Sheviak & W.F. Jennings Habenaria quinqueseta var. macroceratitis (Willdenow) Luer = Habenaria macroceratitis Willdenow Habenaria saccata Greene = Platanthera stricta Lindley Habenaria sparsiflora S. Watson = Platanthera sparsiflora (S. Watson) Schlechter var. sparsiflora Habenaria sparsiflora S. Watson var. laxiflora (Rydberg) Correll = Platanthera sparsiflora (S. Watson) Schlechter Habenaria straminea Fernald = Pseudorchis straminea (Fernald) Soó
128

SYNONYMS AND MISAPPLIED NAMES

Habenaria strictissima Reichenbach f. var. odontopetala (Reichenbach f.) L.O. Williams = Habenaria odontopetala Reichenbach f. Habenaria unalascensis (Sprengel) S. Watson = Piperia unalascensis (Sprengel) Rydberg Habenaria unalascensis (Sprengel) S. Watson var. elata (Jepson) Correll =Piperia elongata Rydberg Habenaria unalascensis (Sprengel) S. Watson var. maritima (Greene) Correll = Piperia elegans (Lindley) Rydberg subsp. elegans Habenaria viridis (Linnaeus) R. Brown in Aiton f. = Coeloglossum viride (Linnaeus) Hartman var. viride Habenaria viridis Linnaeus var. bracteata (Mühlenberg in Willdenow) Reichenbach in Gray = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer Habenaria viridis Linnaeus var. interjecta Fernald = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer Habenaria xandrewsii M. White in G.G. Niles = Platanthera xandrewsii (White) Luer Habenaria xbicolor (Rafinesque) Beckner = Platanthera xbicolor (Rafinesque) Luer Habenaria xcanbyi Ames = Platanthera xcanbyi (Ames) Luer Habenaria xchapmanii (Small) Ames = Platanthera chapmanii (Small) Luer emend. Folsom Habenaria xmedia (Rydberg) Niles = Platanthera huronensis (Nuttall) Lindley Habenella odontopetala (Reichenbach f.) Small = Habenaria odontopetala Reichenbach f. Hammarbya paludosa (Linnaeus) Kuntze = Malaxis paludosa (Linnaeus) Swartz Harrisella filiformis (Swartz) Cogniaux ≠ Harrisella porrecta (Reichenbach f.) Fawcett & Rendle Harrisella porrecta (Reichenbach f.) Fawcett & Rendle = Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten Helleborine gigantea (Douglas) Druce = Epipactis gigantea Douglas in Hooker Hexalectris aphyllus Rafinesque = Hexalectris spicata (Walter) Barnhardt Hexalectris mexicana Greenman = Hexalectris grandiflora (A. Richard & Galeotti) L.O. Williams Hormidium pygmaeum (Hooker) Bentham & Hooker f. = Prosthechea pygmaea (Hooker) W.E. Higgins Ibidium beckii (Lindley) House = Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer Ibidium beckii House = Spiranthes tuberosa Rafinesque Ibidium cernuum (Linnaeus) House = Spiranthes cernua (Linnaeus) L.C. Richard Ibidium floridanum Wherry = Spiranthes floridana (Wherry) Cory emend. P.M. Brown Ibidium gracile (Bigelow) House = Spiranthes lacera (Rafinesque) Rafinesque var. gracilis (Bigelow) Luer Ibidium laciniatum (Small) House = Spiranthes laciniata (Small) Ames Ibidium longilabre (Lindley) House = Spiranthes longilabris Lindley Ibidium lucayanum Britton = Mesadenus lucayanus (Britton) Schlechter Ibidium odoratum (Nuttall) House = Spiranthes odorata (Nuttall) Lindley Ibidium ovale (Lindley) House = Spiranthes ovalis Lindley Ibidium plantagineum (Rafinesque) House = Spiranthes lucida (H.H. Eaton) Ames Ibidium porrifolium Rydberg = Spiranthes porrifolia Lindley Ibidium praecox (Walter) House = Spiranthes praecox (Walter) S. Watson Ibidium strictum (Rydberg) House = Spiranthes romanzoffiana Chamisso Ibidium tortile (Swartz) House = Spiranthes torta (Thunberg) Garay & Sweet Ibidium vernale (Engelmann & A. Gray) House = Spiranthes vernalis Engelmann & A. Gray Isotria affinis (Austin in A. Gray) Rydberg = Isotria medeoloides (Pursh) Rafinesque Lepanthes harrisii Fawcett & Rendell = Lepanthopsis melanantha (Reichenbach f.) Ames Leucorchis albida subsp. straminea (Fernald) A. Löve = Pseudorchis straminea (Fernald) Soó
129

SYNONYMS AND MISAPPLIED NAMES

Limnorchis behringiana Rydberg = Platanthera tipuloides (Linnaeus) Lindley var. behringiana (Rydberg) Hultén Limnorchis chorisiana (Chamisso) J.P. Anderson = Platanthera chorisiana Chamisso Limnorchis convallariaefolius (Fischer) Rydberg = Platanthera convallariifolia Fischer Limnorchis dilatata (Pursh) Rydberg in Britton = Platanthera dilatata (Pursh) Lindley Limnorchis ensifolia Rydberg = Platanthera tescamnis Sheviak & W.F. Jennings Limnorchis hyperborea (Linnaeus) Rydberg = Platanthera hyperborea (Linnaeus) Lindley Limnorchis laxiflora Rydberg = Platanthera sparsiflora (S. Watson) Schlechter Limnorchis leucostachys (Lindley) Rydberg = Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer Limnorchis media Rydberg = Platanthera huronensis (Nuttall) Lindley Limnorchis purpurascens Rydberg = Platanthera purpurascens (Rydberg) Sheviak & W.F. Jennings Limnorchis sparsiflora (S. Watson) Rydberg = Platanthera sparsiflora (S. Watson) Schlechter Limnorchis stricta (Lindley) Rydberg = Platanthera stricta Lindley Limodorum altum Linnaeus = Eulophia alta (Linnaeus) Fawcett & Rendle Limodorum gentianoides Swartz = Triphora gentianoides (Swartz) Ames & Schlechter Limodorum lanceolatum Aublet = Sacoila lanceolata (Aublet) Garay var. lanceolata Limodorum multiflorum (Lindley) Mohr = Calopogon multiflorus Lindley Limodorum pallidum (Chapman) Mohr = Calopogon pallidus Chapman Limodorum parviflorum (Lindley) Nash = Calopogon barbatus (Walter) Ames Limodorum pinetorum Small = Calopogon multiflorus Lindley Limodorum praecox Walter = Spiranthes praecox (Walter) S. Watson Limodorum pulchellum Salisbury = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg Limodorum purpureum Lamark = Bletia purpurea (Lamark) de Candolle Limodorum simpsonii Small = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. simpsonii (Small) Magrath Limodorum tankervilleae Aiton = Phaius tankervilleae (Aiton) Blume Limodorum tuberosum Linnaeus = Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg Liparis nervosa (Thunberg) Lindley ≠ Liparis elata Lindley Listera borealis Morong forma trifolia Lepage = Listera auriculata Wiegand forma trifolia (Lepage) Lepage Listera caurina Piper = Listera banksiana Lindley Listera cordata (Linnaeus) R. Brown subsp. nephrophylla (Rydberg) A. & D. Löve = Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hultén Listera nephrophylla Rydberg = Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hultén Listera reniformis Small = Listera smallii Wiegand Listera retusa Suksdorf = Listera banksiana Lindley Lophiaris carthagenensis (Jacquin) Braem = Trichocentrum carthagenense (Jacquin) M.W. Chase & N.H. Williams Lophiaris lurida (Lindley) Braem = Trichocentrum luridum (Lindley) M.W. Chase & N.H. Williams Lysias hookeriana Rydberg in Britton = Platanthera hookeri (Torrey) Lindley Lysias macrophylla (Goldie) House = Platanthera macrophylla (Goldie) P.M. Brown Lysias menziesii (Lindley) Rydberg = Platanthera orbiculata (Pursh) Lindley Lysias orbiculata Rydberg = Platanthera orbiculata (Pursh) Lindley Lysias orbiculata Rydberg var. pauciflora Jennings = Platanthera orbiculata (Pursh) Lindley forma pauciflora (Jennings) P.M. Brown Lysiella obtusata Rydberg = Platanthera obtusata (Banks in Pursh) Lindley
130

SYNONYMS AND MISAPPLIED NAMES

Malaxis brachypoda (Gray) Fernald forma bifolia (Mousley) Fernald = Malaxis monophyllos (Linnaeus) Swartz forma bifolia Mousley Malaxis ehrenbergii (Reichenbach f.) Kuntze ≠ Malaxis porphyrea (Ridley) Kuntze Malaxis floridana (Chapman) O. Kuntze = Malaxis spicata Swartz Malaxis macrostachys (Lexarza) Kuntze = Malaxis soulei L.O. Williams Malaxis monophyllos (Linnaeus) Swartz var. brachypoda (A. Gray) Morris & Eames = Malaxis brachypoda (Gray) Fernald Malaxis monophyllos (Linnaeus) Swartz var. diphyllos (Chamisso) Luer = Malaxis diphyllos Chamisso Malaxis montana (Englemann) Kuntze = Malaxis soulei L.O. Williams Malaxis tenuis (S. Watson) Ames ≠ Malaxis abieticola Salazar & Soto Arenas Malaxis unifolia var. bayardii nom. nud. = Malaxis bayardii Fernald Malaxis wendtii Salazar ≠ Malaxis porphyrea (Ridley) Kuntze Maxillaria conferta (Grisebach) Schweinfurth in Leon = Maxillaria parviflora (Poeppig & Endlicher) Garay Mesadenus polyanthus (Reichenbach f.) Schlechter ≠ Mesadenus lucayanus (Britton) Schlechter Microstylis brachypoda A. Gray = Malaxis brachypoda (Gray) Fernald Microstylis corymbosa S. Watson = Malaxis corymbosa (S. Watson) Kuntze Microstylis floridana Chapman = Malaxis spicata Swartz Microstylis macrostachys Lexarza = Malaxis soulei L.O. Williams Microstylis monophyllos (Linnaeus) Lindley ≠ Malaxis diphyllos Chamisso Microstylis montana Englemann = Malaxis soulei L.O. Williams Microstylis porphyrea Ridley = Malaxis porphyrea (Ridley) Kuntze Microstylis spicata (Swartz) Lindley = Malaxis spicata Swartz Microstylis tenuis S. Watson in part = Malaxis abieticola Salazar & Soto Arenas Microstylis unifolia (Michaux) Britton, Sterns & Poggenberg = Malaxis unifolia Michaux Neolehmannia difformis (Jacquin) Pabst ≠ Epidendrum floridense Hágsater Neottia auriculata (Wiegand) Szlachetko = Listera auriculata Wiegand Neottia australis (Lindley) Szlachetko = Listera australis Lindley Neottia borealis (Morong) Szlachetko = Listera borealis Morong Neottia calcarata Swartz = Eltroplectris calcarata (Swartz) Garay & Sweet Neottia caurina (Piper) Szlachetko = Listera banksiana Lindley Neottia cinnabarina La Llave & Lexarza = Dichromanthus cinnabarinus (La Llave & Lexarza) Garay Neottia cordata (Linnaeus) Richard = Listera cordata (Linnaeus) R. Brown var. cordata Neottia gracilis Bigelow = Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer Neottia lacera Rafinesque = Spiranthes lacera Rafinesque var. lacera Neottia lucida H.H. Eaton = Spiranthes lucida (H.H. Eaton) Neottia mertensiana (Bongard) Kuntze = Corallorhiza mertensiana Bongard Neottia michuacana La Llave & Lexarza = Dichromanthus michuacanus (La Llave & Lexarza) Salazar & Soto-Arenas Neottia nephrophylla (Rydberg) Szlachetko = Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hultén Neottia ovata (Linnaeus) Bluff & Fingerhut = Listera ovata (Linnaeus) R. Brown Neottia pubescens Willdenow = Goodyera pubescens (Willdenow) R. Brown Neottia smallii (Wiegand) Szlachetko = Listera smallii Wiegand Neottia striata (Lindley) Kuntze = Corallorhiza striata Lindley Oncidium bahamense Nash in Britton & Millspaugh = Tolumnia bahamense (Nash in Britton & Millspaugh) G.J. Braem
131

SYNONYMS AND MISAPPLIED NAMES

Oncidium carthagenense (Jacquin) Swartz = Trichocentrum carthagenense (Jacquin) M.W. Chase & N.H. Williams Oncidium ensatum Lindley ≠ Oncidium floridanum Ames Oncidium luridum Lindley = Trichocentrum luridum (Lindley) M.W. Chase & N.H. Williams Oncidium undulatum (Swartz) Salisbury = Trichocentrum undulatum (Swartz) Ackerman & Chase Oncidium undulatum (Swartz) Salisbury forma flavovirens P.M. Brown = Trichocentrum undulatum (Swartz) Ackerman & M.W. Chase forma flavovirens (P.M. Brown) P.M. Brown Oncidium variegatum (Swartz) ≠ Tolumnia bahamensis (Nash in Britton & Millspaugh) G.J. Braem Ophrys barbata Walter = Calopogon barbatus (Walter) Ames Ophrys cernua Linnaeus = Spiranthes cernua (Linnaeus) L.C. Richard Ophrys cordata Linnaeus = Listera cordata (Linnaeus) R. Brown var. cordata Ophrys ovata Linnaeus = Listera ovata (Linnaeus) R. Brown Ophrys torta Thunberg = Spiranthes torta (Thunberg) Garay & H.R. Sweet Orchis aristata Fischer in Lindley = Dactylorhiza aristata (Fischer in Lindley) Soó Orchis aristata Fischer in Lindley forma perbracteata Lepage = Dactylorhiza aristata var. kodiakensis Luer forma perbracteata (Lepage) P.M. Brown Orchis blephariglottis Willdenow = Platanthera blephariglottis (Willdenow) Lindley Orchis bracteata Mühlenberg in Willdenow = Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer Orchis ciliaris Linnaeus = Platanthera ciliaris (Linnaeus) Lindley Orchis clavellata Michaux = Gymnadeniopsis clavellata (Michaux) Rydberg Orchis conopsea Linnaeus = Gymnadenia conopsea (Linnaeus) R. Brown Orchis cristata Michaux = Platanthera cristata (Michaux) Lindley Orchis dilatata Pursh = Platanthera dilatata (Pursh) Lindley Orchis discolor Pursh = Tipularia discolor (Pursh) Nuttall Orchis flava Linnaeus = Platanthera flava (Linnaeus) Lindley var. flava Orchis grandiflora Bigelow = Platanthera grandiflora (Bigelow) Lindley Orchis hookeriana Oakes = Platanthera hookeri (Torrey) Lindley Orchis huronensis Nuttall = Platanthera huronensis (Nuttall) Lindley Orchis hyperborea Linnaeus = Platanthera hyperborea (Linnaeus) Lindley Orchis integra Nuttall = Gymnadeniopsis integra (Nuttall) Rydberg Orchis lacera Michaux = Platanthera lacera (Michaux) G. Don Orchis latifolius Linnaeus var. junialis Vermeulen = Dactylorhiza majalis (Reichenbach f.) Summerhayes subsp. praetermissa var. junialis (Vermeulen) Senghas*? Orchis leucophaea Nuttall = Platanthera leucophaea (Nuttall) Lindley Orchis nivea Nuttall = Gymnadeniopsis nivea (Nuttall) Rydberg Orchis obtusata Banks in Pursh = Platanthera obtusata (Banks in Pursh) Lindley Orchis orbiculata Pursh = Platanthera orbiculata (Pursh) Lindley Orchis praetermissa Druce = Dactylorhiza majalis (Reichenbach) Summerhayes var. praetermissa (Druce) D.M. Moore & Soó *? Orchis praetermissa Druce var. albiflora Druce = Dactylorhiza majalis (Reichenbach f.) Summerhayes var. praetermissa (Druce) D.M. Moore & Soó forma albiflora (Druce) P.M. Brown Orchis psycodes Linnaeus = Platanthera psycodes (Linnaeus) Lindley Orchis purpurella T. & T.A. Stephenson ≠ Dactylorhiza majalis (Reichenbach f.) Summerhayes subsp. praetermissa (Druce) D.M. Moore & Soó var. junialis (Vermeulen) Senghas Orchis quinqueseta Michaux = Habenaria quinqueseta (Michaux) Eaton Orchis rotundifolia Banks in Pursh = Amerorchis rotundifolia (Banks) Hultén
132

SYNONYMS AND MISAPPLIED NAMES

Orchis rotundifolia (Banks in Pursh) forma angustifolia Rousseau = Amerorchis rotundifolia (Banks) Hultén forma angustifolia (Rousseau) P.M. Brown Orchis rotundifolia (Banks in Pursh) forma beckettiae Boivin = Amerorchis rotundifolia (Banks) Hultén forma beckettiae (Boivin) Hultén Orchis rotundifolia (Banks in Pursh) forma lineata Mousley = Amerorchis rotundifolia (Banks) Hultén forma lineata (Mousley) Hultén Orchis spectabilis Linnaeus = Galearis spectabilis (Linnaeus) Rafinesque Orchis spectabilis Linnaeus forma albiflora Ulke = Galearis spectabilis (Linnaeus) Rydberg forma gordinierii House Orchis spectabilis Linnaeus forma willeyi Seymour = Galearis spectabilis (Linnaeus) Rafinesque forma lilacina (Ames) P.M. Brown Orchis spectabilis Linnaeus var. lilacina Ames = Galearis spectabilis (Linnaeus) Rafinesque forma lilacina (Ames) P.M. Brown Orchis strateumatica Linnaeus = Zeuxine strateumatica (Linnaeus) Schlechter Pelexia cranichoides Grisebach = Cyclopogon cranichoides (Grisebach) Schlechter Pelexia setacea Lindley = Eltroplectris calcarata (Swartz) Garay & Sweet Peramium decipiens (Hooker) Piper = Goodyera oblongifolia Rafinesque Peramium giganteum (Douglas) Salisbury = Epipactis gigantea Douglas in Hooker Peramium ophioides (Fernald) Rydberg = Goodyera repens (Linnaeus) R. Brown forma ophioides (Fernald) P.M. Brown Peramium pubescens (Willdenow) MacMillan = Goodyera pubescens (Willdenow) R. Brown Peramium tesselatum (Loddiges) A. Heller = Goodyera tesselata Loddiges Physurus querceticola Lindley = Platythelys querceticola (Lindley) Garay Physurus sagreanus A. Richard = Platythelys latifolia (Linnaeus ) Garay & Ormerod Piperia dilatata (Pursh) Szlachetko & P. Rutkowski = Platanthera dilatata (Pursh) Lindley var. dilatata Piperia dilatata (Pursh) Szlachetko & P. Rutkowski var. albiflora (Chamisso) Szlachetko & P. Rutkowski =Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour Piperia elegans var. elata (Jepson) Luer = Piperia elongata Rydberg Piperia elongata Rydberg subsp. michaelii (Greene) Ackerman = Piperia michaelii (E. Greene) Rydberg Piperia lancifolia Rydberg = Piperia elongata Rydberg Piperia longispica Durand = Piperia elongata Rydberg Piperia maritima (Greene) Rydberg = Piperia elegans (Lindley) Rydberg subsp. elegans Piperia multiflora Rydberg = Piperia elegans (Lindley) Rydberg subsp. elegans Platanthera albida (Linnaeus) Lindley var. straminea (Fernald) Luer = Pseudorchis straminea (Fernald) Soó Platanthera andrewsii (White) Luer = Platanthera xandrewsii (White) Luer Platanthera blephariglottis (Willdenow) Lindley var. conspicua (Nash) Luer = Platanthera conspicua (Nash) P.M. Brown Platanthera candida (R. Morgan & J. Ackerman) R.M. Bateman nom. illeg. = Piperia candida R. Morgan & J. Ackerman Platanthera clavellata (Michaux) Luer var. clavellata = Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata Platanthera clavellata (Michaux) Luer var. clavellata forma slaughteri P.M. Brown = Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata forma slaughteri (P.M. Brown) P.M. Brown

133

SYNONYMS AND MISAPPLIED NAMES

Platanthera clavellata (Michaux) Luer var. ophioglossoides (Fernald) P.M. Brown = Gymnadeniopsis clavellata (Michaux) Rydberg var. ophioglossoides (Fernald) W.J. Schrenk Platanthera colemanii (Morgan & Glicenstein) R.M. Bateman = Piperia colemanii Morgan & Glicenstein Platanthera cooperi (S. Watson) R.M. Bateman = Piperia cooperi (S. Watson) Rydberg Platanthera dilatata (Pursh) Lindley var. angustifolia Hooker = Platanthera dilatata (Pursh) Lindley Platanthera dilatata (Pursh) Lindley var. chlorantha Hultén = Platanthera huronensis Lindley Platanthera elegans Lindley = Piperia elegans (Lindley) Rydberg Platanthera elongata (Rydberg) R.M. Bateman = Piperia elongata Rydberg Platanthera foetida Geyer in Hooker ≠ Piperia unalascensis (Sprengel) Rydberg Platanthera gracilis Lindley = Platanthera stricta Lindley Platanthera hookeri (Torrey) Lindley var. oblongifolia J.A. Paine = Platanthera hookeri (Torrey) Lindley forma oblongifolia (J.A. Paine) P.M. Brown Platanthera hookeri (Torrey in A. Gray) Lindley var. abbreviata (Fernald) Catling = Platanthera hookeri (Torrey) Lindley forma abbreviata (Fernald) P.M. Brown Platanthera hookeri (Torrey in A. Gray) Lindley var. abbreviata (Fernald) W.J. Schrenk = Platanthera hookeri (Torrey) Lindley forma abbreviata (Fernald) P.M. Brown Platanthera hyperborea (Linnaeus) Lindley ≠ Platanthera aquilonis Sheviak Platanthera hyperborea (Linnaeus) Lindley forma alba M.H.S. Light = Platanthera aquilonis Sheviak forma alba (Light) P.M. Brown Platanthera hyperborea (Linnaeus) Lindley var. huronensis (Nuttall) Luer = Platanthera huronensis (Nuttall) Lindley Platanthera hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Luer = Platanthera purpurascens (Rydberg) Sheviak & W.F. Jennings Platanthera hyperborea (Linnaeus) Lindley var. viridiflora (Chamisso) Kitamura = Platanthera stricta Lindley Platanthera hyperborea (Linnaeus) Lindley var. viridiflora (Chamisso) Luer = Platanthera stricta Lindley Platanthera integra (Nuttall) Luer = Gymnadeniopsis integra (Nuttall) Rydberg Platanthera lacera var. terrae-novae (Fernald) Luer = Platanthera xandrewsii (Niles) Luer Platanthera leptopetala (Rydberg) R.M. Bateman = Piperia leptopetala Rydberg Platanthera leucostachys Lindley = Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer Platanthera menziesii Lindley = Platanthera orbiculata (Pursh) Lindley Platanthera michaelii (E. Greene) R.M. Bateman = Piperia michaelii (E. Greene) Rydberg Platanthera michauxii (Nuttall) Wood = Habenaria simpsonii Small Platanthera nivea (Nuttall) Luer = Gymnadeniopsis nivea (Nuttall) Rydberg Platanthera oligantha Turczaninow = Platanthera obtusata (Banks in Pursh) Lindley subsp. oligantha (Turczaninow) Hultén Platanthera orbiculata var. macrophylla (Goldie) Luer = Platanthera macrophylla (Goldie) P.M. Brown Platanthera parvula Schlechter = Platanthera obtusata (Banks in Pursh) Lindley subsp. oligantha (Turczaninow) Hultén Platanthera repens (Nuttall) Wood = Habenaria repens Nuttall Platanthera saccata (Greene) Hultén = Platanthera stricta Lindley Platanthera sparsiflora (S. Watson) Schlechter var. ensifolia (Rydberg) Luer = Platanthera tescamnis Sheviak & W.F. Jennings Platanthera transversa (Suksdorf) R.M. Bateman = Piperia transversa Suksdorf Platanthera unalascensis (Sprengel) Kurtz = Piperia unalascensis (Sprengel) Rydberg Platanthera unalascensis (Sprengel) Kurtz subsp. elata (Jepson) Taylor & MacBryde = Piperia elongata Rydberg
134

SYNONYMS AND MISAPPLIED NAMES

Platanthera unalascensis (Sprengel) Kurtz subsp. maritima (Greene) de Filipps = Piperia elegans (Lindley) Rydberg subsp. elegans Platanthera yadonii R. Morgan & J. Ackerman) R.M. Bateman = Piperia yadonii R. Morgan & J. Ackerman Platanthera xbicolor (Rafinesque) Luer ≠ Platanthera xlueri P.M. Brown Platanthera xcanbyi (Ames) Luer ≠ Platanthera xbeckneri P.M. Brown Platanthera xchapmanii (Small) Luer = Platanthera chapmanii (Small) Luer emend. Folsom Platanthera xmedia (Rydberg) Luer = Platanthera huronensis (Nuttall) Lindley Platanthera xvossii = XPlatanthopsis vossii (Case) P.M. Brown Platythelys sagreana (A. Richard) Garay = Platythelys latifolia (Linnaeus) Garay & Ormerod Pleurothallis melanantha Reichenbach f. = Lepanthopsis melanantha (Reichenbach f.) Ames Pleurothallis pachyrhachis A. Richard = Bulbophyllum pachyrhachis (A. Richard) Grisebach Poggenberg Pogonia affinis Austin in A. Gray = Isotria medeoloides (Pursh) Rafinesque Pogonia bifaria (Fernald) P.M. Brown & Wunderlin = Cleistes bifaria (Fernald) Catling & Gregg Pogonia cubensis Reichenbach f. = Triphora gentianoides (Swartz) Ames & Schlechter Pogonia divaricata (Linnaeus) R. Brown = Cleistes divaricata (Linnaeus) Ames Pogonia gentianoides (Swartz) Sprengel = Triphora gentianoides (Swartz) Ames & Schlechter Pogonia ophioglossoides (Linnaeus) Ker-Gawler var. brachypogon Fernald = Pogonia ophioglossoides (Linnaeus) Ker-Gawler forma brachypogon (Fernald) P.M. Brown Pogonia pendula (Mühlenberg in Willdenow) Lindley = Triphora trianthophora (Swartz) Rydberg Pogonia trianthophoros (Swartz) Britton, Sterns & Poggenberg = Triphora trianthophoros (Swartz) Rydberg Pogonia trianthophoros Mühlenberg in Willdenow = Triphora trianthophoros (Swartz) Rydberg Pogonia verticillata (Mühlenberg in Willdenow) Nuttall = Isotria verticillata (Mühlenberg in Willdenow) Rafinesque Polyradicion lindenii (Lindley) Garay = Dendrophylax lindenii (Lindley) Bentham in Rolfe Polyrrhiza lindenii (Lindley) Cogniaux = Dendrophylax lindenii (Lindley) Bentham in Rolfe Polystachya flavescens (Lindley) J.K. Small ≠ Polystachya concreta (Jacquin) Garay & Sweet Ponthieva racemosa (Walter) C. Mohr var. brittonae (Ames) Luer = Ponthieva brittoniae Ames Pseudencyclia boothiana (Lindley) V.P. Castro & Chiron = Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Pseudodiphryllum chorisianum (Chamisso) Nevski = Platanthera chorisiana (Chamisso) Reichenbach f. Pseudorchis albida (Linnaeus) Löve & Löve subsp. straminea (Fernald) Löve & Löve = Pseudorchis straminea (Fernald) Soó Sacoila lanceolata (Aublet) Garay var. squamulosa (Kunth) Szlachetko = Sacoila squamulosa (Kunth) Garay Satyrium adnatum Swartz = Pelexia adnata (Swartz) Sprengel Satyrium elatum Swartz = Cyclopogon elatus (Swartz) Schlechter Satyrium latifolium Linnaeus = Platythelys latifolia (Linnaeus) Garay & Ormerod Satyrium repens Linnaeus = Goodyera repens (Linnaeus) R. Brown Scaphyglottis parviflora Poeppig & Endlicher = Maxillaria parviflora (Poeppig & Endlicher) Garay Schiedeella confusa (Garay) Espejo & López-Ferrari = Deiregyne confusa Garay Schiedeella fauci-sanguinea (Dod) Burns-Balogh ≠ Schiedeella arizonica P.M. Brown Schiedeella parasitica (A. Richard & Galeotti) Schlechter ≠ Schiedeella arizonica P.M. Brown Schiedeella rubrocallosa (Robinson & Greenman) Burns-Balogh = Microthelys rubrocallosa (Robinson & Greenman) Garay
135

SYNONYMS AND MISAPPLIED NAMES

Serapias austinae (A. Gray) A.A. Eaton = Cephalanthera austiniae (A. Gray) Heller Serapias helleborine Linnaeus = Epipactis helleborine (Linnaeus) Cranz* Serapias latifolia Hudson (rank unspecified) atrorubens Hoffman = Epipactis atrorubens (Hoffman) Besser Serapias polystachya Swartz = Tropidia polystachya (Swartz) Ames Spiranthes adnata (Swartz) Bentham in Fawcett = Pelexia adnata (Swartz) Sprengel Spiranthes beckii Lindley = Spiranthes lacera (Rafinesque) Rafinesque var. gracilis (Bigelow) Luer Spiranthes brevifolia Chapman = Spiranthes longilabris Lindley Spiranthes brevilabris Lindley var. floridana (Wherry) Luer = Spiranthes floridana (Wherry) Cory emend. P.M. Brown Spiranthes cernua (Linnaeus) L.C. Richard var. ochroleuca (Rydberg) Ames = Spiranthes ochroleuca (Rydberg) Rydberg Spiranthes cernua (Linnaeus) L.C. Richard var. odorata (Nuttall) Correll = Spiranthes odorata (Nuttall) Lindley Spiranthes cinnabarina (La Llave & Lexarza) Hemsley = Dichromanthus cinnabarinus (La Llave & Lexarza) Garay Spiranthes confusa (Garay) Kartesz & Gandhi = Deiregyne confusa Garay Spiranthes costaricensis Reichenbach f. = Beloglottis costaricensis (Reichenbach f.) Schlechter Spiranthes cranichoides (Grisebach) Cogniaux = Cyclopogon cranichoides (Grisebach) Schlechter Spiranthes decipiens Hooker = Goodyera oblongifolia Rafinesque Spiranthes durangensis Ames & Schweinfurth ≠ Deiregyne confusa Garay Spiranthes elata (Swartz) L.C. Richard = Cyclopogon elatus (Swartz) Schlechter Spiranthes fauci-sanguinea Dod ≠ Schiedeella arizonica P.M. Brown Spiranthes gracilis (Bigelow) Beck = Spiranthes lacera (Rafinesque) Rafinesque var. gracilis (Bigelow) Luer Spiranthes gracilis (Bigelow) Beck var. brevilabris (Lindley) Correll = Spiranthes brevilabris Lindley Spiranthes gracilis (Bigelow) Beck var. floridana (Wherry) Correll = Spiranthes floridana (Wherry) Cory emend. P.M. Brown Spiranthes graminea Lindley ≠ Spiranthes delitescens Sheviak Spiranthes grayi Ames = Spiranthes tuberosa Rafinesque Spiranthes intermedia Ames ≠ Spiranthes casei Catling & Cruise var. casei Spiranthes lanceolata (Aublet) Leon = Sacoila lanceolata (Aublet) Garay var. lanceolata Spiranthes lanceolata (Aublet) Leon var. paludicola Luer = Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Sauleda, Wunderlin & Hansen Spiranthes lucayana (Britton) Cogniaux = Mesadenus lucayanus (Britton) Schlechter Spiranthes michuacana (La Llave & Lexarza) Hemsley = Dichromanthus michuacanus (La Llave & Lexarza) Salazar & Soto-Arenas Spiranthes orchioides (Swartz) A. Richard = Sacoila lanceolata (Aublet) Garay Spiranthes parasitica A. Richard & Galeotti ≠ Schiedeella arizonica P.M. Brown Spiranthes parviflora (Chapman) Ames = Spiranthes ovalis Lindley Spiranthes plantaginea Rafinesque = Spiranthes lucida (H.H. Eaton) Ames Spiranthes polyantha Reichenbach ≠ Mesadenus lucayanus (Britton) Schlechter Spiranthes romanzoffiana Chamisso var. diluvialis (Sheviak) S.L. Welsh = Spiranthes diluvialis Sheviak Spiranthes romanzoffiana Chamisso var. porrifolia (Lindley) Ames & Correll = Spiranthes porrifolia Lindley Spiranthes rubrocallosa Robinson & Greenman = Microthelys rubrocallosa (Robinson & Greenman) Garay Spiranthes squamulosa (Kunth) Leon = Sacoila squamulosa (Kunth) Garay
136

SYNONYMS AND MISAPPLIED NAMES

Spiranthes stricta (Rydberg) A. Nelson in J.M. Coulter & A. Nelson = Spiranthes romanzoffiana Chamisso Spiranthes tortilis (Swartz) Richard = Spiranthes torta (Thunberg) Garay & Sweet Spiranthes tuberosa var. grayi (Ames) Fernald = Spiranthes tuberosa Rafinesque Spiranthes unalascensis Sprengel = Piperia unalascensis (Sprengel) Rydberg Spiranthes vernalis Engelmann & Gray ≠ Spiranthes casei Catling & Cruise Spiranthes xaustralis P.M. Brown nom. illeg. = Spiranthes xmeridionalis P.M. Brown Spiranthes xsteigeri Correll = Spiranthes ochroleuca (Rydberg) Rydberg Stelis gelida (Lindley) Pridgeon & M.W. Chase = Pleurothallis gelida Lindley Stenorrhynchos cinnabarina (La Llave & Lexarza) Lindley = Dichromanthus cinnabarinus (La Llave & Lexarza) Garay Stenorrhynchos cinnabarinum (La Llave & Lexarza) Lindley var. paludicola (Luer) W. J. Schrenk = Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Sauleda, Wunderlin, and B.F. Hansen Stenorrhynchos lanceolatum (Aublet) Richard in Sprengel = Sacoila lanceolata (Aublet) Garay var. lanceolata Stenorrhynchos michuacanum (La Llave & Lexarza) Lindley = Dichromanthus michuacanus (La Llave & Lexarza) Salazar & Soto-Arenas Stenorrhynchos orchoides (Swartz) L.C. Richard = Sacoila lanceolata (Aublet) Garay Stenorrhynchos squamulosum (Kunth) Sprengel = Sacoila squamulosa (Kunth) Garay Tamayorkis wendtii (Salazar) R. González & Szlachetko = Malaxis wendtii Salazar Tipularia unifolia Britton, Sterns & Poggenberg = Tipularia discolor (Pursh) Nuttall Tolumnia variegata (Swartz) G.J. Braem ≠ Tolumnia bahamensis (Nash in Britton & Millspaugh) G.J. Braem Trichocentrum maculatum (Aublet) M. Chase & N.A. Williams nom. illeg. = Trichocentrum undulatum (Swartz) Ackerman & Chase Triorchis beckii House = Spiranthes tuberosa Rafinesque Triorchis cernua (Linnaeus) Nieuwland = Spiranthes cernua (Linnaeus) L.C. Richard Triorchis cernua Nieuwland var. ochroleuca Farwell = Spiranthes ochroleuca (Rydberg) Rydberg Triorchis gracile (Bigelow) Nieuwland = Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer Triorchis grayi (Ames) Nieuwland = Spiranthes tuberosa Rafinesque Triorchis laciniata House = Spiranthes laciniata (Small) Ames Triorchis longilabris House = Spiranthes longilabris Lindley Triorchis ochroleuca Nieuwland = Spiranthes ochroleuca (Rydberg) Rydberg Triorchis odorata (Nuttall) Nieuwland = Spiranthes odorata (Nuttall) Lindley Triorchis ovalis Nieuwland = Spiranthes ovalis Lindley Triorchis plantaginea Nieuwland = Spiranthes lucida (H.H. Eaton) Ames Triorchis praecox (Walter) Nieuwland = Spiranthes praecox (Walter) S. Watson Triorchis romanzoffiana (Chamisso) Nieuwland = Spiranthes romanzoffiana Chamisso Triorchis stricta (Rydberg) Nieuwland = Spiranthes romanzoffiana Chamisso Triorchis vernalis House = Spiranthes vernalis Engelmann & A. Gray Triphora cubensis (Reichenbach f.) Ames = Triphora gentianoides (Swartz) Ames & Schlechter Triphora latifolia G. Luer = Triphora amazonica Schlechter Triphora pendula (Mühlenberg in Willdenow) Nuttall = Triphora trianthophoros (Swartz) Rydberg Triphora yucatanensis Ames ≠ Triphora rickettii Luer Tropidia eatonii Ames = Tropidia polystachya (Swartz) Ames Vanilla articulata Northrop = Vanilla barbellata Reichenbach f. Vanilla eggersii Rolfe ≠Vanilla dilloniana Correll
137

SYNONYMS AND MISAPPLIED NAMES

Vanilla fragrans (Salisbury) Ames = Vanilla planifolia Andrews Vanilla mexicana Miller = Vanilla inodora Schiede
Note: The International Plant Name Index remarks that in Triphora trianthophora the "-os" ending in "trianthophoros" (Greek) denotes feminine gender and has corrected all citations of trianthophora to trianthophoros.

LITERATURE CITED:

Brown, P.M. and S.N. Folsom. 2003. The Wild Orchids of North America, North of Mexico. Gainesville: University Press of Florida. _____. 2004. Wild Orchids of the Southeastern United States. Gainesville: University Press of Florida. _____. 2005. Wild Orchids of Florida expanded and updated. Gainesville: University Press of Florida. _____. 2006a. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region. Gainesville: University Press of Florida. _____. 2006b. Wild Orchids of the Pacific Northwest and Canadian Rockies. Gainesville: University Press of Florida. _____. 2006c. Wild Orchids of the Prairies and Great Plains Region of North America. Gainesville: University Press of Florida. _____. 2007. Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey. Gainesville: University Press of Florida. _____. 2008. Field Guide to the Wild Orchids of Texas. Gainesville: University Press of Florida. Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford: Oxford University Press, Volume 26 (2002) including the Orchidaceae. International Plant Names Index. http://www.ipni.org/index.html Luer, C.A. 1972. The Native Orchids of Florida. Bronx: New York Botanical Garden. _____. 1975. The Native Orchids of the United States and Canada Excluding Florida. Bronx: New York Botanical Garden.

ACKNOWLEDGEMENTS:

Although many people have contributed information and advice the author specifically thanks Chuck Sheviak, Scott Stewart, Lucy Dueck, and Jim Fowler for their corrections and suggestions. Compiled by Paul Martin Brown April 2008

A NEW COMBINATION
In 1900 Oakes Ames described a new color variant of Orchis spectabilis with the lip solid lilac in color as are the petals and sepals. Subsequently Frank Seymour did the same in 1970 as Orchis spectabilis forma willeyi with the labellum being pink instead of white. These are clearly the same color form, as the plants vary from lilac to pink but in both cases the lip is other than white and colored the same as the rest of the perianth. The Ames publication takes precedent and a new combination and status is needed. Ames did list the form in his Enumeration of the Orchids of the U.S. & Canada in 1924. Galearis spectabilis (L.) Raf. forma lilacina (Ames) P.M. Brown stat. & comb. nov. Basionym: Orchis spectabilis (L.) Raf. var. lilacina Ames Amer. Gardening 21: 375. 1900 Synonyms: Orchis spectabilis forma willeyi F. Seym. Rhodora 72: 48. 1970 Galearis spectabilis forma willeyi (F. Seym.) P.M. Brown Wild Flower Notes 3(1): 16. 1988

Special thanks to Tom & Jackie Nelson, Carolyn Beans (HUH), and Elizabeth Allen (VT) for their assistance.

138

Bransilver: MY FAVORITE THINGS

MY FAVORITE THINGS Connie Bransilver
Connie is best known to native orchid enthusiast for her book Wild Love Affair, with dramatic photographs of south Florida‘s orchids.

Beeswarm orchid, Cyrtopodium punctatum

139

Bransilver: MY FAVORITE THINGS

Clamshell orchid, Prosthechea cochleata var. triandra

140

Bransilver: MY FAVORITE THINGS

crested coralroot Hexalectris spicata

141

Bransilver: MY FAVORITE THINGS

ghost orchid Dendrophylax lindenii

all photographs © Connie Bransilver Barrow

142

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII

THE SPIRANTHES FORMERLY KNOWN AS PARKSII
Lucy A. Dueck When fellow Spiranthes aficionado Scott Stewart walked me out of the auditorium after my talk to a group of native orchid enthusiasts on the molecular phylogeny of North American Spiranthes a couple years ago, he exclaimed that I had just rewritten the book on the genus. Scott was particularly interested because we had provided evidence for genetic distinction between the two state endangered or proposed-endangered species he worked on, S. brevilabris and S. florida, which some had confused with each other. But the most surprising result we found was about a federally listed species endemic only to Texas, S. parksii, which apparently has an identity crisis. It was at that moment that Scott coined the term ―the Spiranthes formerly known as parksii‖, reminiscent of a famous musician. With some help from the American Orchid Society, I had been working for several years on this research project to unravel Spiranthes relationships using genetics, and was joined by Dr. Ken Cameron, then-director of the molecular systematics program at the New York Botanical Garden (now at the University of Wisconsin–Madison). Until recently, our work has only been published in the form of an extended abstract on the conservation genetics of the entire genus (Dueck and Cameron, 2007). But we knew that we needed to focus a second paper specifically on S. parksii, given our discovery, because its protection status was coming up for review soon. Such reviews are required through the Endangered Species Act for plants and animals listed as federally threatened or endangered. The U.S. Fish & Wildlife Service is charged with using the best data available to assess the listee‘s recovery progress and decide if the species should be removed or re-classified. Determining whether the listee is really a ‗good species‘ is not usually an issue at this stage, but obviously is quite important for continued protection if its identity is questionable. We therefore are publishing a peer-reviewed technical paper in the journal Conservation Genetics about S. parksii‘s species status based on genetic data to provide information for this decision (Dueck and Cameron, 2008). Below I present a synopsis of our work from that paper, along with a few additional comments, for those readers who may not have access to the journal. Spiranthes parksii, or Navasota ladies‘-tresses, is a fall-blooming terrestrial orchid found in a dozen counties of east central Texas and in one county near the Louisiana border. It was discovered and described in the late 1940s but disappeared for many years thereafter, only to be re-discovered about thirty years later. Shortly afterward, in 1982, it was declared an endangered species and as such has influenced land use decisions where it occurs ever since (Correll, 1947; Luer, 1975; Catling and McIntosh, 1979; Liggio and Liggio, 1999). However, S. parksii is among 15 Spiranthes found in Texas and was initially confused with two of them that bloom nearby about the same time – S. lacera var. gracilis and S. cernua. But S. parksii has several characteristics that differentiate it morphologically from other ladies‘-tresses such as smaller slightly gaping flowers, longer upturned sepals, and white tips on its floral bracts, although there is certainly variation within (Figure 1). It does share some other important features with the common S. cernua though – the same duplicate chromosome number that indicates they are both polyploids,
143

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII

and the same asexual type of multi-embryo seed development that essentially produces clones. Its cousin S. cernua is not without variation within it also – it comes in at least two forms in Texas, open and closed (Figure 2). The genus Spiranthes has long been known as incredibly difficult to sort out due to variation and hybridization. To complicate matters further, S. cernua (with duplicate sets of chromosomes) is considered to be the recipient of gene flow from several other closely related Spiranthes (with single sets of chromosomes), thereby broadly extending its adaptability and range across North America. It could thus be called a compilospecies, and the entire group of Spiranthes involved is called the ―cernua complex‖. Spiranthes parksii is thought to also be a member of the complex, but more as an outcome, like fellow polyploid S. cernua, than a contributor (Sheviak, 1982; 1991). In fact, Sheviak and Brown (2002) suspected that S. parksii may actually be some variant of S. cernua, attributing its unusual morphology to at least partial peloria (an aberration from the normal floral symmetry which in orchids means the petals and the lip, normally quite different, begin to look like each other).

Figure 1. Three forms of Spiranthes parksii: beautiful like a beeswax candle, typical, and contorted.

Molecular phylogenetics is the study of how a set of organisms are related. It uses a variety of genetic tools at different levels of discrimination, and it provides a phylogeny, or ‗family tree‘, that is inferred from the information and used to show the relationships within the lineage in order to hypothesize the pattern and process of the lineage‘s evolution. Different molecular markers, and even different regions within the same kind of markers, can provide different genealogical trees that may not be exactly the same. But when they are in general agreement, it strengthens the case for the truth about the relationships. A graduate student at Texas A&M University, Cathy Walters, using a fine-scale set of molecular markers (AFLPs and microsatellites), was looking for population differences among colonies of Spiranthes parksii. But
144

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII

instead she discovered there was basically no difference between the endangered species and the open form of its local common cousin S. cernua when she compared them; however, she did find they both were a little different from the closed form of S. cernua (Walters 2005). This happened at about the same time we also made the same discovery within our broader scale project, only we were using sequences of DNA regions appropriate for a somewhat larger scale at the genus level. So that is when we decided to take a closer look at more Spiranthes samples specifically from Texas.

Figure 2. Open, half-open, and closed forms of Spiranthes cernua as found in Texas.

We used 60 samples from nine Spiranthes species (11 each from S. parksii and Texas S. cernua; S. lacera var. gracilis, S. longilabris, S. magnicamporum, S. praecox, S. vernalis, and S. sylvatica), plus a relative in the same subtribe as Spiranthes (Sacoila lanceolata var. lanceolata) to provide perspective in the genetic analyses. These samples were mostly from Texas, but some specimens of the same species collected in other parts of the U.S. were included, again for perspective. The plant tissue was dried until DNA could be extracted from it (or DNA already extracted was received, using the same S. parksii samples as Walters). Then I employed PCR to amplify the extracted DNA with four different pairs of primers. This means that a specific chunk of DNA was replicated many times. The four primer pairs I used represented segments of DNA in all three plant genomes – one nuclear segment, two chloroplast segments, and one mitochondrial segment. The nuclear genome is inherited from both parents (pollen and seed), and the latter two genomes are inherited only from the mother (seed). It is important to get the full picture by looking at data from both types of genetic inheritance to see if they differ. The amplified DNA segments were subsequently sequenced, and every sequence was checked for correctness. This is where Ken took over and analyzed the sets of sequences in special software to produce a tree for each of
145

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII

the four DNA segments plus a fifth tree from combining all data; each tree and its branching pattern was then statistically tested. Oftentimes the combined data tree provides more statistical confidence in the way it branches than the single trees do. Our analyses generated over 3000 base pairs (A, C, G, or T and its complement in double-stranded DNA), each pair being an individual bit of information that could differ among the samples to tell them apart. But even before Ken performed any special data analysis, I manually reviewed those from Spiranthes parksii and when compared to the open form of S. cernua from Texas, they were exactly the same in every base pair for every DNA segment I sequenced. Even the closed form of S. cernua from Texas had only three differences in two of its samples from either S. parksii or open S. cernua. Keep in mind that the DNA segments that show essentially no difference here are the same ones that can tell other species of Spiranthes apart – that is why it is important to include many samples and species from the genus for comparison. When Ken analyzed the sequence data, he found that one of the chloroplast DNA segments (cp3) was very good at grouping the samples into like ‗clades‘, and it even contained an easy identifier, an extra 25 base pairs of DNA, present only in the group containing Spiranthes parksii. This ‗parksii group‘ also contained all S. cernua from Texas, as well as S. cernua from Nebraska and from an unusual escarpment site in South Carolina, but did not contain S. cernua from several other eastern sites (although they were ‗sisters‘ – very closely related). In turn, the ‗parksii group‘ and ‗cernua sisters‘ combined with most other species in a somewhat amorphous group that was sister to a very strong group containing S. praecox and S. sylvatica. The tree shape from the other chloroplast DNA segment (cpL) was very similar to the cp3 tree, although relationships among the non-cernua-parksii/praecox-sylvatica species were less clear. Mitochondrial sequences (mt7) did not provide much definitive information. But the tree from the nuclear DNA segment (nr) combined the ‗parksii group‘ and ‗cernua sisters‘ together (except for the Nebraska sample), which grouped with all other species except S. praecox-sylvatica and S. lacera var. gracilis. So from this biparentally inherited DNA segment, there were three strong groups – the latter two and all other species, with S. lacera var. gracilis at the base of the tree, farthest away from S. parksii. Finally, the tree from combining sequences of all four DNA segments showed a branching pattern very similar to that of the cp3 tree above only stronger, particularly with a separate ‗parksii group‘. Figure 3 shows a simplified version of this tree. All of the original trees are available in our technical publication at www.springerlink.com. So what does all this mean? Well, not only are the sequences of Spiranthes parksii and Texas S. cernua the same, but also our analyses position S. parksii broadly within the single group of all S. cernua. This group cannot be split apart for S. parksii without splintering it apart into many taxa below the level of species. There are several other important similarities as well between the two species, besides the same duplicate chromosome number and asexual development of multi-embryo seeds mentioned earlier – they can be found growing as close as inches to each other, they bloom at the same time, and they cannot be told apart from leaves alone (Wilson, 2002). Granted, the flowers of S. parksii look somewhat different than those of S. cernua, but S. cernua from Texas (particularly the closed form) looks different than S. cernua from eastern states. It is likely that both S. parksii and closed-form S. cernua are peloric mutants, a condition not uncommon within an orchid species and which could explain their differing morphology. For all the above reasons, we suggest that S. parksii be considered synonymous, or the same as, S. cernua in the broad sense of the species. There are those who would like to consider other issues concerning the status of Spiranthes parksii. One of those issues is whether it is a hybrid; certainly its duplicate chromosome number and lack of intermediacy between the only two living possibilities – S. cernua and S. lacera var. gracilis – rule out simple hybrid status. Spiranthes parksii is a polyploid though (meaning it has
146

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII

more than the usual two sets of chromosomes), which can occur either from a ―permanent‖ hybridization event between species or as a duplicating of chromosome number within a species. But a combination of our and Walters‘ molecular evidence and its asexual reproduction suggest no hybridization was involved. In the S. cernua compilospecies, however, chromosome duplication is rampant. According to Tate et al. (2005), different episodes of genome duplication involving the same species may result in different phenotypes having the same genotype – that is, organisms that do not look alike but have the same genetic makeup. Differential gene expression among versions of Spiranthes cernua may also have the potential to change how that version develops and looks, and could be induced by several factors. Polyploidy is one such factor, along with epigenetics – heritable molecular modifications from several possible mechanisms that do not change a DNA sequence (Soltis et al., 2003; Mauricio, 2005; Rapp and Wendel, 2005). Another possible factor is infection by different mycobionts – mycorrhizal fungi essential for initial establishment of wild orchid seeds. Some researchers have found differences in gene regulation or floral variation within terrestrial orchid species depending on which fungus was present (Watkinson, 2002; Taylor and Bruns, 1999; Taylor et al., 2003). However, all of these factors are admittedly speculative long shots to explain morphological differences, and none of them can turn S. parksii into a different species with a separate genotype from S. cernua. Thus according to the phylogenetic species concept, which defines a species in terms of being classified as the smallest possible group through evolution (Mayden, 1997), Spiranthes parksii cannot be separated from the S. cernua group. Even if an attempt to subdivide the S. cernua group into many varieties and forms was made, it would be an impossible task that would not benefit the conservation of orchids overall (Pillon and Chase, 2007). Therefore, we suggested in our paper that S. parksii is merely another of the many (unofficial) forms taken by the complex species S. cernua. As for the fate of its listing as an endangered species, it will be up to the U.S. Fish & Wildlife Service to make that decision very soon.
REFERENCES Catling, P.M. and K.L. McIntosh. 1979. Rediscovery of Spiranthes parksii Correll. Sida 8: 188-93. Correll, D.S. 1947. A new Spiranthes from Texas. American Orchid Society Bulletin 16: 400. Dueck, L.A. and K.M. Cameron. 2007. Sequencing re-defines Spiranthes relationships, with implications for rare and endangered taxa. Lankesteriana 7(1-2): 190-95. _____. 2008. Molecular evidence on the species status and phylogenetic relationships of Spiranthes parksii, an endangered orchid from Texas. Available via OnlineFirst at: http://www.springerlink.com, DOI 10.1007/s10592-007-9501-1; Conservation Genetics (in press). Liggio, J. and A.O. Liggio. 1999. Wild Orchids of Texas. University of Texas Press, Austin, Tex. Luer, C.A. 1975. The Native Orchids of the United States and Canada excluding Florida. New York Botanical Garden, Bronx N.Y.. Mauricio, R. 2005. The ‗bricolage‘ of the genome elucidated through evolutionary genomics. New Phytologist 168: 1-4. Mayden, R.L. 1997. A hierarchy of species concepts: the denouement in the saga of the species problem. In: Claridge, M.F., H.A. Dawah, and M.R. Wilson (eds.) Species: The Units of Biodiversity, Chapman & Hall, London, pp. 381-424. Pillon, Y. and M.W. Chase. 2007. Taxonomic exaggeration and its effects on orchid conservation. Conservation Biology 21(1): 263-65. Rapp, R.A. and J.F. Wendel. 2005. Epigenetics and plant evolution. New Phytologist 168: 81-91. Sheviak, C.J. 1982. Biosystematic study of the Spiranthes cernua complex. Bulletin No. 448, New York State Museum, Albany, N.Y. _____. 1991. Morphological variation in the compilospecies Spiranthes cernua (L.) L.C. Rich.: ecologically-limited effects of gene flow. Lindleyana 6: 228-34. Sheviak, C.J. and P.M. Brown. 2002. Orchidaceae, Vol. 26, Spiranthes. Flora of North America. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=131021. 147

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII Soltis, D.E., P.S. Soltis, and J.A. Tate. 2003. Advances in the study of polyploidy since Plant Speciation. New Phytolologist 161: 173-91. Tate, J.A., D.E. Soltis, and P.S. Soltis. 2005. Polyploidy in plants. In: Gregory, T.R. (ed.) The Evolution of the Genome, Elsevier, Burlington, Mass. pp. 371-426. Taylor, D.L., T.D. Bruns, J.R. Leake, and D.J. Read. 2002. Mycorrhizal specificity and function in mycoheterotrophic plants. In: van der Heijden, M.G.A., and I.R. Sanders (eds.) Mycorrhizal Ecology, Ecological Studies, vol. 157. Springer-Verlag, Berlin, pp. 375-413. Taylor, D.L., T.D. Bruns, T.M. Szaro, and S.A. Hodges. 2003. Divergence in mycorrhizal specialization within Hexalectris spicata (Orchidaceae), a nonphotosynthetic desert orchid. American Journal of Botany 90: 1168-79. Walters, C. 2005. Genetic relationships among Spiranthes parksii and congeneric species. M.Sc. thesis, Texas A&M University, College Station, Tex. Watkinson, J.I. 2002. Characterization of two genes, trehalose-6-phosphate synthase /phosphatase and nucleotide binding protein, shown to be differentially regulated in roots of Cypripedium parviflorum var. pubescens grown with a mycorrhizal fungus Thanatephorus pennatus. Ph.D. dissertation, Virginia Polytechnic Institute & State University, Blacksburg, Va. Wilson, H.D. 2002. Distinctive features of S. parksii as compared to other fall-blooming species in the Navasota Flora. http://www.csdl.tamu.edu/FLORA/hdwsp/parksii_id_1001a.htm. ACKNOWLEDGEMENTS The author thanks the U.S. Fish & Wildlife Service for permit #TE100419-0 to transport samples, and the SC Department of Natural Resources for permission to collect some samples. Special gratitude goes out to Ken Cameron for being my accomplice and reviewer, Cathy Walters for sharing DNA and information, Ryan Hammons for showing me examples, Jim Fowler and Scott Stewart for helpful suggestions, and to the sample collectors: S. Bentley, R. Bischof, P.M. Brown, J. Fowler, F. Galloway, J. Liggio, S. Shriver, S. Stewart, and B. Summers. Materials for this project were funded by a grant in 2004-5 from the American Orchid Society, and facilities plus some time were supported by the University of Georgia Research Foundation through award #DE-FC09-07SR22506 by the U.S. Department of Energy. All photos by Lucy Dueck. Lucy A. Dueck, Savannah River Ecology Lab, Aiken, SC 29802; current address: P.O. Box 1413, Athens, TX 75751; email: lucydueck@msn.com. GLOSSARY ancestral node – a junction on a ‗family tree‘ representing a speciation event where the ancestral lineage splits into at least two descendant lineages, based on changes in quantifiable traits used to reconstruct the tree. base pair – two nucleotides (chemical compounds that are structural units) on complementary DNA strands connected by hydrogen bonds; A (adenine) and T (thymine) are always paired, and C (cytosine) and G (guanine) are always paired. clade – a taxonomic group of organisms including a single ancestor and all of its descendants; i.e., a branch and its tips of a ‗family tree‘. compilospecies – a model or concept used to describe a conglomerate species (usually polyploid) that evolved and expanded its range by hybridizing with more geographically restricted but closely related species (usually diploid); all species involved are often called a ‗complex‘. differential gene expression – different cell types or activities resulting from genetically identical genes being activated differentially at several possible levels in the development of an organism, potentially producing different phenotypes. molecular marker – a genetic trait specific to a particular organism or group of organisms that can be revealed by any of a number of laboratory and analytical methods, the choice of which method depending on the level of specificity required. monophyletic – evolved from a single common ancestor (―one line‖). phylogenetic species concept – one of many ways to define what constitutes a species, and in which a species is identified as the smallest biological entity that is diagnosable and monophyletic, resulting from natural selection and descent. polyploidy – the presence of three or more chromosome sets in an organism, resulting from the multiplication of one chromosome set, known as autopolyploidy, or from the merger of structurally different chromosome sets, called allopolyploidy (Tate et al. 2005). resolved/unresolved (re nodes in ‗family tree‘) – A resolved node is a junction where an ancestral lineage splits into two immediate descendant lineages, while an unresolved node will have more than two immediate descendant

148

Dueck: THE SPIRANTHES FORMERLY KNOWN AS PARKSII lineages. Unresolved nodes are called polytomies, and most typically are interpreted as uncertainty about relationships, usually because we lack information about branching order for the taxa within it. For further explanation, consult websites such as: http://evolution.berkeley.edu/evolibrary/article/0_0_0/phylogenetics_01

149

McGrath: PLATANTHERA PALLIDA

CONTRIBUTIONS TO THE STATUS AND MORPHOLOGY OF PLATANTHERA PALLIDA, PALE FRINGED ORCHIS
Reprinted with permission from the Long Island Botanical Society Newsletter 18:1 2008.

Robert T. McGrath
Author’s Note: After reading the excellent book review prepared by Eric Lamont in the summer issue (vol. 17, 2007) of the LIBS newsletter on the Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey by Paul Martin Brown, I immediately went on-line to purchase my own copy. When it arrived a few days later I too was extremely impressed with the attention to detail, wide range of supplementary material, photography, and beautiful illustrations. Of particular interest for me was Brown‘s recognition of two populations of Platanthera located in eastern Long Island as warranting endemic species status. These populations have long been considered to be Platanthera cristata, the yellow-crested orchid. Naturalists and botanists have long noted that the two eastern Long Island populations, located at Napeague and the Walking Dunes at Montauk differ from other populations of P. cristata because of their pale yellow coloration as opposed to the deeper orange normally found in the species. Brown first described these colonies as a valid species, Platanthera pallida in 1992 and he continues to describe them as endemic only to eastern Long Island from two sizable colonies in this latest publication (2007). I was especially intrigued by this designation as I had studied these populations during the summers of 1982 and 1983 and found morphological aberrations not previously reported in the literature at the time. Being a young undergraduate full of vim and vigor and jumping from one research project to the next, I never completed preparing my findings for publication. The paper presented here represents an original accounting of the observations that I made at that time.

OBSERVATIONS ON AN ABERRANT POPULATION OF THE YELLOWCRESTED ORCHID, PLATANTHERA CRISTATA

Original Manuscript
INTRODUCTION During the first week of August, 1983 four populations of the crested yellow orchid, Platanthera cristata (Michx.) Lindl., were located in Suffolk County, Long Island, New York. Each of the populations appeared to correspond morphologically to the description given by Correll (1950) for Habenaria cristata (Michx.) R. Br., and by Luer (1975) for Platanthera cristata, but contained, either partially or entirely, individuals that exhibited a light cream-yellow color similar to the variation noted by Carpenter (1959). A closer study of flowers collected from one of the populations revealed that it contained numerous individuals with supernumerary anthers and fusion of one or both of the lateral petals with the dorsal sepal (Fig. 1). Teratological and floral abnormalities are seemingly rare in Platanthera. The only published report is that of an instance of carpellody, the transformation of a stamen to a pistil, in Platanthera bifolia (Rytz, 1921).

150

McGrath: PLATANTHERA PALLIDA

DISTRIBUTION AND HABITAT Platanthera cristata is a coastal plain species inhabiting moist pine woodland clearings from eastern Texas northeast to Massachusetts and south along the Atlantic Coast to Florida. The four Long Island populations were located in similar habitats. A description of each population and its habitat follows:\ 1. Oakdale – Approximately ten plants, first located on 9 August 1982 along a fire lane (these plants were mowed prior to blooming in 1983) in Pine Barrens habitat. These individuals were found growing in moist sandy loam in open sunlight, in association with Platanthera blephariglottis (white fringed orchid), Drosera rotundifolia (roundleaf sundew), Carex sp. (sedge), and Clethra alnifolia (sweet pepper bush). All of these orchid flowers exhibited a pale canary-yellow color. 2. Sag Harbor – Approximately 60 plants first seen on 5 August 1983. These plants were found growing in moist sandy loam along a roadside ditch in full sun, and under a dense tree canopy of Pinus rigida (pitch pine) in association with Rhexia virginica (meadow beauty), Bartonia virginica (yellow screwstem), Carex sp. (sedge), Polygala polygama (racemed milkwort), and (under dense tree canopy) Kalmia angustifolia (sheep laurel), Comptonia peregrina (sweet fern), Myrica pensylvanica (bayberry), and Ilex glabra (inkberry). The perianths of the individuals in this population exhibited two distinct color variations, a deep yellow-orange and a light cream-yellow. Both color variations existed along the road as well as under the dense tree canopy, with no distinguishable preference. 3. Napeague – Approximately 1000 plants first seen on 5 August 1983 and then re-visited on 12 August 1983. These individuals were found growing in damp sandy loam under a stand of Pinus rigida in association with Cypripedium acaule (pink lady's slipper). All individuals exhibited a light cream-yellow color. 4. Montauk – Approximately 500 plants first seen on 12 August 1983. All of the plants were found growing in dry sandy loam under a stand of Pinus rigida and Quercus alba (white oak), and in association with Arctostaphylos uva-ursi (bearberry), Cypripedium acaule, and Panicum sp. (panic grass). Flowers of all of the individuals were pale creamy-yellow in color. All four populations were previously known, and had been documented by earlier collectors. Specimens deposited at the New York Botanical Garden by Roy Latham and labeled Platanthera cristata (Latham 6992, 6993) contain population descriptions and localities very similar to the Napeague and Montauk populations described above. Another locality documented by Latham (Latham 7308, 7490 at NYBG) is described as follows: East Hampton, Long Island, colony of one hundred plants, in Pine Barrens Swamp. On 7 Aug 1888, Arthur K. Harrison collected P. cristata from "Sag Harbor" and noted, "1 plant with orange flowers, 2 plants with lemon flowers" (voucher specimen at Harvard Univ. Herbaria; Lamont, personal communication, 2007). On 1 Aug 1951, Roy Latham, C.K. Brooks & H.F. Dunbar collected P. cristata from "dry barrens near Montauk Point", and noted "flowers lemon yellow;? x H. blephariglottis (which is not near)". On 10 Aug 1948, C.S. Bryan collected it from "Montauk Point", and noted "abnormal in the flower color which is cream-color or light sulphurous yellow" (Voucher specimens of both latter collections housed at Harvard Univ. Herbaria; Lamont, personal communication, 2007).

151

McGrath: PLATANTHERA PALLIDA

METHODS AND OBSERVATIONS Flowers from the three populations observed blooming in August 1983 were collected by randomly walking through each colony and removing flowers from various plants. Specimens were fixed in 45% ethyl alcohol, 45% distilled water, 5% glacial acetic acid, and 5% formalin. There was no attempt made to correlate plant characters and flowers during collection. Measurements at 75X magnification were made of the spur length, dorsal and lateral sepal width and length, petal length and width, and lip length, and these were found to be compatible with the measurements reported by Correll (1950). These results are shown in Table 1.
Characteristics Lip width Dorsal sepal length Dorsal sepal width Lateral petal length Lateral petal width Lateral sepal length Lateral sepal width Spur length * when present Correll Range Sag Harbor Napeague Montauk

---3-5 2-3 2-4 2-4
3-4

2 4 2.5 3 3
3

2 4 4 3* 2.5*
3

2 4 2.4 3 3
4

2-3 5-10

3 6

2.75 6

2.5 ----

Table 1. Measurements (mm.) of floral characters in Long Island populations of Platanthera cristata.

As described by both Correll and Luer, the dorsal sepal of Platanthera cristata is characteristically elliptic to suborbicular, with a slight notch at the obtuse apex. During analysis of flowers collected from the Napeague colony, however, numerous individuals were noted which showed fringing on either the right, left, or both sides of the dorsal sepal (Fig. 1).

Figure 1. Diagrammatic line drawings of dissected flowers from the Napeague colony showing various forms of dorsal sepal fringing.

In all instances where fringing occurred, the lateral petal on the fringed side was absent. Furthermore, a thickened tissue region running the vertical length of the dorsal sepal, approximately 0.5 mm from the fringed edge was observed. Of the 72 flowers collected from the Napeague colony, 11 displayed sepal fringing on both sides and had no lateral petals; ten displayed sepal fringing on the right side and had no right lateral petal; eight displayed sepal fringing on the left side and had no left lateral petal; and 42 displayed a typical part arrangement for Platanthera cristata. These observations lead to the conclusion that the aberrant fringing on the dorsal sepal is the result of the developmental fusion of the dorsal sepal with one or both of the lateral petals.
152

McGrath: PLATANTHERA PALLIDA

In addition to aberrations on the dorsal sepal, at least four flowers were noted having supernumerary anthers arising on the column (Fig. 2). All anthers appeared to be sexually functional and pollinia were produced in all the anther sacs. Examination of various pollinia revealed a club-shaped mass of mealy pollen, caudicle, and viscidium typical for Platanthera. Supernumerary anther sacs varied from four to eight per flower. DISCUSSION: The lack of data on measurements of non-floral characters, on the cytology of the aberrant floral parts, on pollination success, and on the heritability of aberrant characters, prohibits determining the significance and nature of these findings. While the aberrations may have arisen through hybridization of Platanthera cristata with another member of yellow orchid complex, this is not likely in the author‘s opinion. The occurrence in general of hybrids between P. cristata and either P. ciliaris (L.) Lindl. or P. blephariglottis (Willd.) Lindl. is widely accepted (e.g. Folsom, 1979; Luer, 1975; Correll, 1950; Ames, 1908; Small,1903). Both of the latter species do occur on Long Island, but P. ciliaris is quite rare and is known from only a few localities, while P. blephariglottis is widely scattered in small colonies of no more than 60 plants. The hybrid Platanthera ciliaris x P. cristata was originally named as a species, Blephariglottis chapmanii Small (1903); was later treated as a hybrid Platanthera xchapmanii (Small) Luer (see Luer, 1975); and was most recently again treated as a species, Platanthera chapmanii (Small) Luer by Folsom (1979). Folsom revealed that the morphology of this cross is intermediate in most respects between that of P. ciliaris and P. cristata. However, as noted previously, flowers collected from all three Long Island colonies correspond morphologically to the description of P. cristata. Furthermore, the pollinating system for P. cristata would seem to be sufficiently different from P. ciliaris to prohibit cross pollination of the two species. As noted by Folsom (1979) and Smith and Snow (1976), pollination of P. ciliaris is effected primarily by butterflies of the family Papilionadae, the swallowtails, while bumble bees (Bombus sp.) are the primary pollinators of P. cristata. During my own investigations, a bumble bee was observed visiting approximately eight plants in the Napeague colony during a 30-second interval at approximately 7:00 PM on 12 August 1983. This individual was not captured, and while it can be considered only a potential pollinator, its role as a pollinator of P. cristata would seem likely given this observation and those made by Folsom (1979). In summary, evidence from structural characteristics, from pollination and from local scarcity all lead to rejecting P. ciliaris as a likely parent. Though the hybrid Platanthera blephariglottis x cristata is more plausible based on the greater availability of P. blephariglottis, it would have to be disregarded for the morphological and geographic reasons given previously, and the fact that pollination in P. blephariglottis is effected primarily by night flying moths (Smith and Snow, 1976). From my field observations, the Napeague colony does not appear to grow in a habitat that is drastically different from the other populations. Environmental factors such as soil acidity, nutrient content, and moisture were not recorded, however, and may play a role in the occurrence of these aberrations. In his study on abnormal flowers, Meyer (1966) noted the importance of the external environment in promoting metamorphosis of floral organs. Wherry, in his exhaustive studies on soil acidity and its effects on floral distribution (1918, 1920, 1927), suggested optimum soil acidity values for many orchids; but their significance and effect, if any, on plant morphology were not stated. That a relationship exists between aberrant dorsal sepals and supernumerary anthers is unlikely, as all three variations of fringing, as well as the normal flower condition, were found on flowers with more than the ordinary number of anthers. Because of the random collection of the
153

McGrath: PLATANTHERA PALLIDA

studied flowers, it is impossible to say whether the aberrations are genetic mutations, though Meyer (1966) concluded that non-heritable abnormalities are unlikely to be recorded unless they are produced by a stable physiological or environmental manipulating factor. CONCLUSIONS, LITERATURE UPDATES, AND PRESERVATION STATUS OF EXISTING POPULATIONS

2007 UPDATE

Although clearly differing in color from populations occurring elsewhere on Long Island, the colonies studied on eastern Long Island were considered to be Platanthera cristata up to and including the time when I first studied them in the summers of 1982 and 1983 (Lamont et al., 1988). In his original publication Brown (1992) concluded that the morphological differences, namely, the descending, recurved lips and very short spurs, that these individuals display warranted their being designated a separate species. The findings presented here, while not directly supporting Brown‘s claims, do indeed support the evidence that the eastern populations of this orchid, now formally referred to as Platanthera pallida P. M. Br. (1992), do in fact demonstrate certain distinctly different features from other populations of Platanthera cristata. Just how distinct they are, however, continues to be a matter of speculation. Brown (2007) recognized that Sheviak (2002) considers P. pallida to be a cline of P. cristata, meaning that while the plants described as P. pallida demonstrate changes in characteristics of the typical P. cristata, these changes are part of a broader set of changes which occur over a geographic area. Sheviak specifically noted that, while plants described as P. pallida are not sufficiently distinct to warrant separate species status from P. cristata, they are also not merely hybrids. Sheviak based this conclusion on the fact that many of the features upon which Brown based his designation overlap both species (Sheviak, 2002). He further stated that he has obtained chromosome counts of P. pallida of 2n=42, the same as P. cristata and the other species in the group. This would rule out the possibility that the colonies are allopolyploids, i.e. polyploids with chromosomes derived from different species (Sheviak, personal communication, 2007). I believe the fusion between floral parts and the production of supernumerary anthers described here lend credence to Sheviak‘s claim. That is, that the colonies described by Brown as Platanthera pallida are the product of a partially stabilized introgression, meaning they are the backcrossing of hybrids of two separate plant populations, the progeny of which are producing new genes into a wild population (Sheviak, 2002). This seems to be a far more plausible designation in my mind, especially given the fact that the populations in question could not be isolated from other populations of P. cristata for more than 5000 to 8000 years given the geologic history of Long Island and especially the region in which the colonies in question exist. It is unfortunate that the flower samples that were collected in 1983 have almost all been destroyed. The specimen displayed in Fig. 2 is one of the few remaining samples still preserved. In an attempt to learn whether or not the observations made in 1983 could be documented in the current population I revisited the colony in Napeague, referred to now by many as the Lazy Point orchids, on 11 August 2007. Racemes were removed from approximately 25 plants and fixed in 45% ethyl alcohol, 45% distilled water, 5% glacial acetic acid, and 5% Bio-Gard. Figs. 3 and 4 demonstrate fringing on the left (Fig. 3) and both (Fig. 4) sides of the dorsal sepal with the corresponding lateral petal(s) absent. No instances of supernumerary anthers were observed in the samples collected in 2007. The population located in Oakdale was revisited on 12 August 2007, along a fire lane known locally as Cordwood Road, in Connetquot River State Park. Four
154

McGrath: PLATANTHERA PALLIDA

plants were found and all displayed a light creamy-yellow coloration. (Platanthera blephariglottis, which had been found in 1982 and 1983, could not be re-located during the 2007 visit.) Fig. 5 depicts a flower collected from this population, clearly showing a typical P. cristata with distinctly separate lateral petal and longer lip than those found in populations further east.

2

3

4

5

Figure 2. Flower of P. cristata showing multiple (supernumerary) anther sacs. Pollinia were typical for P. cristata. Figure 3. Flower showing fringing on the left side of the dorsal sepal and the presence of the right lateral petal. Figure 4. Flower showing fringing on both the left and right side of the dorsal sepal and the absence of both lateral petals.Figure 5. Flower of P. cristata taken from population located along Cordwood Road in Oakdale. Of particular note is the distinctly separate lateral petal and longer lip.

Of critical importance to discussions of the evolutionary status of the Long Island populations of Platanthera is the overall protection that each population so desperately needs. Of particular concern is the severe denuding that the Lazy Point (Napeague) population is suffering
155

McGrath: PLATANTHERA PALLIDA

because of deer browsing. During the visit made on 11 August 2007, the population was estimated at 2000 to 3000 plants. While this is quite large, it is estimated that 7 out of every 10 plants in some spots were browsed. This is much more than has ever been observed in previous years. What long-term impact deer browsing will have on the population is uncertain, but considering the potential that these plants have for evolutionary study it would be a shame if they were to be diminished in abundance or, even worse, extirpated because of the overpopulation of white-tailed deer that Long Island is experiencing. Acknowledgements I want to thank Eric Lamont for encouraging me to revisit the work I began 24 years ago and for his constructive criticism of the manuscript; and Charles Sheviak for his manuscript review and for his data on the genetics of Platanthera pallida.
LITERATURE CITED Ames, O. 1908. Notes on Habenaria. Rhodora 10: 70-71. Brown, P. M. 1992. Platanthera pallida (Orchidaceae), a new species of fringed orchis from Long Island, New York, U. S. A. Novon 2:308-11. Brown, P.M. and S.N. Folsom. 2007. Wild Orchids of the Northeast: New England, New York, Pennsylvania and New Jersey. University Press of Florida, Gainesville. Carpenter, G. M. 1959. A Color variant of Habenaria cristata. Bulletin of the Torrey Botanical Club 86: 137-38. Correll, D. S. 1950. Native Orchids of North America. Chronica Botanica Company, Waltham, Mass. Folsom, J. P. 1979. The True Nature of the Putative Natural Hybrid Platanthera x chapmanii (Orchidaceae). M.S. Thesis, Vanderbilt University, Nashville, Tenn. Lamont, E. E., J. M. Beitel & R. E. Zaremba. 1988. Current status of orchids on Long Island, New York. Bulletin of the Torrey Botanical Club 115: 113-21. Luer, C. A. 1975. The Native Orchids of the United States and Canada excluding Florida. The New York Botanical Garden, Bronx, New York. Meyer, V. G. 1966. Flower abnormalities. Botanical Review (Lancaster) 32: 165-218. Rytz, W. 1921. Blütenanomalien. Sitzungsber. Bern. Bot. Ges. 7 Okt. In Mitth. Naturf. Ges. Bern 1921: xxv-xxvi. Sheviak, C. J. 2002. Platanthera, pp. 511-71. In Flora of North America Editorial Committee (editors), Magnoliophyta: Liliidae: Liliales and Orchidales. Flora of North America North of Mexico. vol. 26. Oxford University Press, Oxford. Small, J. K. 1903. Flora of the Southeastern United States. Published by the author, New York. Smith, G. R. & G. E. Snow. 1976. Pollination ecology of Platanthera (Habenaria) ciliaris and P. blephariglottis (Orchidaceae). Botanical Gazette 137: 133-40. Wherry, E. T. 1918. The reaction of the soils supporting the growth of certain native orchids. Journal of the Washington Academy of Science 8: 589-98. ______. 1920. Soil acidity--its nature, measurement, and relationship to plant distribution. Report (Annual) of the Board Regents of the Smithsonian Institution. 2622: 247-68. ______. 1927. The soil reaction of saprophytic orchids. Journal of the Washington Academy of Science 17: 35-38. Robert T. McGrath, Department of Science and Technology Education, Northport- East Northport Union Free School District, Northport, New York, 11768 Email: rmcgrath@northport.k12.ny.us

156

McGrath: PLATANTHERA PALLIDA

Coming in July….. ADDITIONAL DISTRIBUTIONAL INFORMATION REGARDING SPIRANTHES STELLATA WILD ORCHIDS IN THE 21ST CENTURY: RECENT PUBLICATIONS TWO (NOT-SO) NEW COLOR FORMS FROM THE CENTRAL APPALACHIANS SPIRANTHES CASEI IN THE NORTHEASTERN UNITED STATES THE GENUS SACOILA IN THE UNITED STATES
AND MUCH MORE….

157

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

1

PLATANTHERA PALLIDA

158

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

PLATANTHERA PALLIDA - FIFTEEN YEARS OF COMPARISONS
Paul Martin Brown When I first saw plants of the pale-flowered Platanthera cristata (Michaux) Lindley growing in eastern Long Island, New York, in 1989 I admittedly had seen very few populations of the species but knew that these distinctive plants were noticeably different. On that day we had just been guided to a small population of typical P. cristata, P. blephariglottis (Willdenow) Lindley, and several excellent plants of P. xcanbyi (Ames) Luer (P. blephariglottis x P. cristata). Having the opportunity to see all of these taxa in the span of a few hours provided the basis for comparisons. Repeated visits in subsequent years to additional eastern Long Island sites, as well as viewing plants growing in New Jersey and Massachusetts only reinforced my original opinion as to the uniqueness of the Montauk peninsula plants. In preparation for publication of Platanthera pallida P.M. Brown as a new species (Brown, 1992) 327 herbarium specimens and several hundred living plants in a variety of habitats were examined and special attention was paid to those described as ‗pale yellow‘ ‗yellow‘ or ‗lemon colored‘. For the most part these pale-colored plants of P. cristata, designated as P. cristata forma straminea P.M. Brown, occurred as scattered individuals in populations of typical orange-colored P. cristata. Only in a few instances were plants of P. xcanbyi found. In the ensuing years many more plants of P. cristata and P. xcanbyi (and within the southeastern coastal plain P. xbeckneri P.M. Brown (2002) [P. cristata x P. conspicua]) were examined both in the herbarium and in situ. Quite simply put, none of the above taxa have the distinctive combination of characters bolded below in a modification of the original description of P. pallida. Plants of Platanthera pallida grow from (20-)29 to 65(-84) cm tall, are glabrous and distinctly glaucous; the 2-3 lower leaves are sheathed, strongly keeled, and conduplicate, growing to 25(-30) cm long and 3(-5) cm wide when flattened; the upper leaves are reduced to 3 to 5 linear bracts below the inflorescence. The racemose inflorescence of (18-)24 to 80(-112) flowers is (5-)10 to 20(-27) cm long and 2.5 to 4.0 cm in diameter and densely flowered except in very tall individuals; the lower floral bracts are usually equal to or often exceeding the pedicellate ovary and decreasing in length upwards, about 2.2 cm long. The perianth is very pale orange-yellow in bud opening to pale cream, the petals and lip often aging to a deeper creamy yellow or fading to a dull white; the lateral sepals are 3 mm long by 3 mm wide, reflexed, widest at the middle and tapering to a rounded apex and truncated base; the dorsal sepal is 3 mm long by 3 mm wide, concave, entire and arching forward to partially enclose the fringed petals; the petals are obovate, 3.25 mm long by 2 mm wide, fringed at the tip, with the fimbriae usually forked and clearly visible to the sides of the dorsal sepal; the lip is ligulate, 3 mm long by 2 mm wide (exclusive of fringe); recurved or, rarely, descending, the margin with forked fimbriae up to 3 mm long, with the fimbriae near the base perpendicular to the lip; the spur is 5--6 mm long, tubular, strongly curved, and either obtuse or bulbous at the tip; the

159

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

nectary orifice is T-shaped; the column, with the viscidia, is 2.5--3 mm apart, and the fruit a capsule, 1.0 to 1.6 cm long.

Figure 2. Platanthera pallida P.M. Brown. --- A. Habit - B. Perianth; front/side view - C. Petal – D. Lip - E. Column; side view - F. Dorsal Sepal - G. Lateral Sepal Drawn by Stan Folsom

160

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

KEY TO THE ORANGE, YELLOW, AND WHITE-FRINGED ORCHIDS OF NORTH AMERICA 1a. Spur less than 10 mm long; shorter than ovary...2 1b. Spur greater then 10 mm long; exceeding ovary...3 2a. Lip recurved, lateral sepals reflexed, dorsal sepal entire, spur 5--6 mm long, obtuse; perianth cream-colored...Platanthera pallida (Figs. 1-4) 2b. Lip projecting forward, lateral sepals porrect, dorsal sepal emarginate, spur 7-8 mm long, acute; perianth orange to yellow...Platanthera cristata (Fig. 5) i. Flowers pale yellow… Platanthera cristata forma straminea (Fig. 6) 3a. Flowers white…4 3b. Flowers otherwise…5 4a. Lip narrowed to a slender isthmus (1:3) at the base i. Lip margin delicately fringed; plants restricted to the southeastern and Gulf coastal plains…P. conspicua (Fig. 10) ii. Lip margin merely erose; plants of the Cumberland Plateau…P. integrilabia 4b. Lip with a broadened isthmus (1:1) at the base; plants widespread in eastern North America…P. blephariglottis (Fig. 7) 5a. Spur 10-15 mm long, exceeding ovary; flowers may be white, cream, pale yellow, yellow, or orange i. Plants intermediate in morphology between P. blephariglottis and P. cristata; isthmus of lip 1:1…Platanthera xcanbyi (Fig. 8) ii. Plants intermediate in morphology between P. conspicua and P. cristata; isthmus of lip 1:3…Platanthera xbeckneri (Fig. 9) 5b. Spur greater than 15 mm long; greatly exceeding the ovary…6 6a. Flowers bright orange or yellow…P. ciliaris (Fig. 11) 6b. Flowers may be cream, pale yellow, yellow, coffee-colored, or bicolored; raceme usually 3 cm or more in diameter i. Plants intermediate in morphology between P. blephariglottis and P. ciliaris; isthmus of lip 1:1…Platanthera xbicolor (Fig. 12) ii. Plants intermediate in morphology between P. conspicua and P. ciliaris; isthmus of lip 1:3…Platanthera xlueri (Fig. 13) Couplets 5 and 6 have attempted to provide a key to the hybrids in this group. Unfortunately it is not always as clear-cut as this. Perhaps it is best said that hybrids usually occur as random individuals within or adjacent to colonies of either or both parents. Observing plants that appear to be intermediate in color and morphology is the first clue to detecting hybrids. Color is highly variable as one colony with Platanthera xcanbyi on Long Island has, in a single season, provided hybrid plants from deep orange to pure white! Because the ranges of P. blephariglottis and P. conspicua have virtually no overlap, although they are approximate in one area of eastern North Carolina, the respective hybrids with P. cristata are also well separated geographically. The relative size and shape of the isthmus at the base of the lip on the whiteflowering species is always evident in the hybrids. The hybrid between P. ciliaris and P. cristata, P. xchannellii has a spur that is equal in length to the ovary and lip and although color does not aid in identifying this hybrid as the plants of
161

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

both parents and hybrids are shades of orange, the size of the raceme in P. xchannellii is intermediate between that of the parents (Brown & Folsom, 2008). It is very important to remember that any description of Platanthera cristata prior to 1992, and those after that date that do not recognize P. pallida or are based upon earlier descriptions, may include some of the same characters that so typify P. pallida, i.e. short spur, recurved lip, etc. This problem is an often overlooked aspect of taxonomy, especially by the local native enthusiast, and may lead to unneeded confusion when comparing the ‗old‘ broadly defined species and the ‗new‘ narrowly defined species. During the past twenty-five years many new species have been described that were segregated from more widespread and familiar species. If the morphological descriptions of those traditional species are still relied upon they usually are found to contain criteria that are distinctive to the more recently described species, e. g. Platanthera pallida within P. cristata. Some excellent examples are cited below. These recently published or revalidated species should be compared with descriptions in Fernald (1950), Correll (1950), Gleason (1978), Gleason and Cronquist (1991) and, although to a lesser extent, Luer (1972, 1975). It is important to remember that contributions to the Orchidaceae in Flora of North America north of Mexico (2002) do not necessarily reflect research through that date. At best, the research that influenced species treatments concluded in 2000 with minor revisions in 2001. Many of the treatments were completed up to 10 years earlier and were not substantially revised. Calopogon oklahomensis D.H. Goldman 1995 from Calopogon tuberosus (Linnaeus) BSP/C. barbatus (Walter) Ames Cypripedium kentuckiense C.F. Reed 1981 from Cypripedium parviflorum Salisbury complex Epidendrum floridense Hágsater 1993 from Epidendrum difforme Jacquin Galeandra bicarinata G.A. Romero & P.M. Brown 2000 from Galeandra beyrichii Reichenbach f. Platanthera aquilonis Sheviak 1999 from Platanthera hyperborea (Linnaeus) Lindley Platanthera conspicua (Nash) P.M. Brown revalidated 2004 from Platanthera blephariglottis (Willdenow) Lindley (originally published as Habenaria conspicua Nash) Platanthera praeclara Sheviak & M.L. Bowles 1986 from Platanthera leucophaea (Nuttall) Lindley Platanthera purpurascens (Rydberg) Sheviak & W.F. Jennings revalidated 1997 from Platanthera hyperborea var. purpurascens (Rydberg) Luer originally published as Limnorchis purpurascens Rydberg in 1901 Platanthera tescamnis Sheviak & W.F. Jennings 2006 from Platanthera sparsiflora (S. Watson) Schlechter Platanthera zothecina (L.C. Higgins & S.L. Welsh) Gandhi & Kartesz 1986 from Platanthera sparsiflora (S. Watson) Schlechter Spiranthes magnicamporum Sheviak 1973 from Spiranthes cernua (Linnaeus) L.C. Richard Spiranthes ochroleuca (Rydberg) Rydberg and S. odorata (Nuttall) Lindley were both described as species much earlier but merged within S. cernua and not revalidated until 1976 and 1980 Spiranthes stellata P.M. Brown, L.A. Dueck & K.M. Cameron 2008 from Spiranthes romanzoffiana Chamisso Spiranthes sylvatica P.M. Brown 2002 from Spiranthes praecox (Walter) S. Watson The following discussion is taken from my original paper published in Novon 2: 308-311, 1992 and is included here for those who do not have easy access to that publication.

162

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

Growing among Pinus rigida in dry interdunal hollows on eastern Long Island, New York, is a Platanthera with small flowers, short spur and superficial resemblance to P. cristata. The most obvious difference at first encounter is the uniform pale cream color of the small flowers. Unlike typical P. cristata in the north, the plants are locally abundant. Detailed examination and observations of P. cristata throughout its range, including all known populations from Long Island, have been made to see if they reveal morphological differences. Measurements were taken of the critical characters, i.e. petals, sepals, lip, spur, cilia and column, of 327 herbarium specimens and 128 living specimens, including 78 plants growing on Long Island. Numerous photographs and drawings, published and unpublished, were also reviewed (Rickett, 1966). Special attention was given to those designated as 'light yellow or pale' in coloration. With the exception of the plants in question all specimens and living material examined fell well within the criteria for typical P. cristata (Correll, 1950; Luer, 1975). Herbarium specimens and living plants of the northern hybrids, P. xcanbyi, P. xbicolor, and P. ciliaris x P. blephariglottis, (reported from Michigan) were also examined to see if they might be similar to the Long Island plants. They were found to be distinct in all respects. Platanthera pallida shows insufficient features to assume its parentage is the same as P. xcanbyi. Although the small flower size and pale coloration can be found in many plants of P. xcanbyi the longer spur so characteristic of that hybrid is lacking in P. pallida. The reflexed sepals and recurved lip which are critical features of P. pallida are also present in P. blephariglottis. Comparison with plants from Michigan appearing to be the cross between P. ciliaris and P. blephariglottis showed no similarities, as the putative hybrids are much larger than P. pallida and the spur much longer, as it is in both parents. One of the remarkable aspects of P. pallida is the uniformity of its floral morphology. All the critical floral characters, i.e. perianth dimensions, color, positioning of floral parts etc., have little, if any, variation throughout all populations. Platanthera pallida occurs as three populations in two sites in the Town of East Hampton, eastern Long Island. The plants appear to have been first discovered by Roy Latham in 1926 (Latham, 1940). By 1948 and subsequent years the stations were visited by several botanists and orchid enthusiasts (Lamont, Beitel, and Zaremba, 1988). Latham's initial site near Montauk supports two distinct current populations. They are separated by nearly 1/4 mile of duneland. In each of these populations the plants are widespread and somewhat scattered, but retain their habitat preference. Adjacent to the areas that support Platanthera pallida are numerous swales and bogs. Typical P. cristata, if present, would be found in these wetter areas, as it is in the Pinelands of New Jersey, a region of similar topography. Careful searches have revealed no other species of Platanthera in either the immediate area or for some miles around. The other population is located west of Napeague Harbor. Several specimens collected by Latham (NYS) in 1928-29 and simply labeled "Napeague" may be from this site. It was not until 1975 that G.E. Lotowycz found the current site and collected her first specimen - as P. cristata - from there. In contrast to the Montauk site, where the plants are widely scattered, here P. pallida is concentrated in a much smaller area and in larger numbers. Again there are adjacent swales and small bogs, but no other Platanthera species to be found. Companion plants are essentially the same in both locales. Platanthera pallida is consistently observed to be restricted to the oldest, most stable Pinus rigida stands within the dunes.
163

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

An interesting side note is that all the flowers on any given Platanthera pallida inflorescence always set seed. This may account for the large, but local, colonies. Also the nearxeric habitat of P. pallida is so unlike that of P. cristata found elsewhere in the Northeast. Platanthera cristata colonies observed in Massachusetts, New York (Long Island), New Jersey, and Delaware are all in considerable moister habitats with companion plants more typical of coastal Atlantic white cedar, Chamaecyparis thyoides, wetlands and open trailside ditches often with Drosera filiformis, Xyris spp., Lachnanthes caroliniana, Lycopodiella spp., Pogonia ophioglossoides and Platanthera blephariglottis. The lack of these typical moisture loving species in the P. pallida sites is notable, although many of these and additional wetland species are found nearby in moister interdunal hollows, but never plants of P. pallida. After fifteen years of reviewing and searching for more colonies of Platanthera pallida and plants that indicate any intermediacy with other species of Platanthera, especially in southern New England, Long Island, and coastal New Jersey, the only populations remain those near Montauk, Long Island, New York. These plants are currently treated at the species level in several publications (Brown and Folsom, 1997; 2007; Chapman, 1997; Lamont, 1996; McGrath, 2008). Regardless if one chooses to recognize them at the species level or as a cline within P. cristata (Sheviak, 2002) they remain a group of distinctive and unique plants among the orchids of North America.

ACKNOWLEDGEMENTS: I wish to especially thank Eric Lamont for suggesting that I write a fifteen-year update on Platanthera pallida and his comments and suggestions on the current manuscript. Scott Stewart, Skip Blanchard, and Margaret Conover also provided comments and suggestions. Observations over the years from many friends have contributed to or reinforced information in this paper; for those observations I am most grateful. LITERATURE CITED: Brown, P.M. 1992. Platanthera pallida (Orchidaceae), a new species of fringed orchis from Long Island, New York, U. S. A. Novon 2: 308-311. ______. 2002. Revalidation of Platanthera conspicua. North American Native Orchid Journal 8: 3-14. Brown, P.M. and S.N. Folsom. 1997. Wild Orchids of the Northeastern United States. Cornell University Press. Ithaca. ______. 2007. Wild Orchids of the Northeast: New England, New York, Pennsylvania and New Jersey. University Press of Florida, Gainesville. ______. 2008 Field Guide to the Wild Orchids of Texas. University Press of Florida, Gainesville. Chapman, W. 1997. Orchids of the Northeast. Syracuse University Press. Syracuse. Correll, D. S. 1950. Native orchids of North America north of Mexico. Chronica Botanica, Waltham, Mass. Fernald, M.L. 1950. Gray’s Manual of Botany 8th edition. American Book Company. Cambridge, Mass. Gleason, H.A. 1978. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. 3 volumes. Hafner Press. New York. Gleason, H.A. and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd edition. New York Botanical Garden. New York. Lamont, E.E. 1996. Atlas of the orchids of Long Island, New York. Bulletin of the Torrey Botanical Club 123: 157-166. Lamont, E.E., J.M. Beitel and R.E. Zaremba. 1988. Current status of orchids on Long Island, New York. Bulletin of the Torrey Botanical Club. 115: 113--121. Latham, R.A. 1940. Distribution of wild orchids on Long Island. Long Island Forum. 3: 103--107. Luer, C.A. 1972. Native orchids of Florida. New York Botanical Garden. Bronx, New York. ______. 1975. Native orchids of the United States and Canada (excluding Florida). New York Botanical Garden. Bronx, New York. McGrath, R. 2008. Contributions to the status and morphology of Platanthera pallida, Pale Fringed Orchis. Long Island Botanical Society Newsletter 18:1-6. Rickett, H.W. 1966. Wild Flowers of the United States 1:1. New York Botanical Garden/ McGraw-Hill, New York. Sheviak, C.J. 2002. Platanthera, pp. 511-571. In Flora of North America Editorial Committee (editors), Magnoliophyta: Liliidae: Liliales and Orchidales. Flora of North America North of Mexico. vol. 26. Oxford University Press, Oxford.

164

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

3 Platanthera pallida

4

5

5 Platanthera cristata
165

6 forma straminea

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

center: Platanthera xcanbyi right: Platanthera blephariglottis

8 7

7 8

5 Platanthera cristata

center: Platanthera xbeckneri right: Platanthera conspicua

9

10

6

166

Brown: PLATANTHERA PALLIDA 15 YEARS OF COMPARISONS

center: Platanthera xbicolor right:: Platanthera blephariglottis

12

8

11 Platanthera ciliaris

center: Platanthera xlueri right: Platanthera conspicua

13

10

167

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS
Reprinted with permission from McAllen International Orchid Society Journal 9(3) 9-16. 2008

Robert J. Ferry

Preface Past research by this worker concentrated on the members of the tropical genus Stanhopea, with only scattered samples from terrestrial orchids. For this line of research, it was fruitful to develop and employ a method of obtaining a casting of the leaf surface for certain epiphytes, but for terrestrial orchids in particular, there is really no substitute for working with live fresh leaf material. Specimens undergo deformation when dried, and although morphological studies may be productive to a certain extent when they are reconstituted, critical cell measurements remain compromised. The leaf casting system developed by this worker for his doctoral research is limited when studying decidedly pubescent to rugose surfaces, ones clad with a heavy waxy cuticle, or for leaves of very light texture as seen in some terrestrial orchid species. For these plants, with their thinner leaves, a small sample of fresh, living material is not only the best avenue, but-in fact-the only one that will provide realistic results. For practical purposes, the shipment of such material to a microscopic center is out of the question. Delays in transit are bound to degrade both the material and any results obtained. The microscopic and photographic equipment must be brought relatively close to the collection area to ensure the highest quality of data obtained photographically. This sounds complicated, but, in reality, is relatively simple. One simply packs equipment, travels to a location a short distance from the collection site, sets up equipment, and then collects, and photographs. The photographic record then becomes an enduring data sample that may not only be filed with a particular archival repository, but electronically shared worldwide, freeing archival centers with having to pack and mail material. While leaf material may be retained in a herbarium, it becomes relatively useless from a morphological standpoint. Both anatomical and life cycle information need to be provided to field biologists nationally and internationally. To do this, plant specimens and seed must be allowed to be collected for investigations into the roles played by various soil fungi in the life cycles of different terrestrial orchid species. As a practical matter, this means collecting seed, flasking it using various solutions, and determining which soil fungus (or fungi) plays the optimum role in plantlet survival. In this field, seminal work has already begun (Zettler, 1997a, 1997b; Stewart & Zettler, 2002; Stewart, Zettler & Minso, 2003; Stewart & Kane, 2006, and others). However, if this work is to be expanded to the profit of field biologists globally, restrictions on collecting and propagating need to be removed for qualified workers to conduct serious studies. Such work also implies keeping a detailed photographic and text record of the life cycle of each species, for dissemination not only to domestic governmental field biologists, but particularly to those field biologists in countries where certain terrestrial orchids are at serious risk of being harvested (many illegally!) to extinction! None of these activities are suggested as commercial enterprises! They are suggested to learn more about these endangered plants, and provide field workers with detailed anatomical and life cycle information in order to aid in the recognition at all life cycle
168

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

stages, and assist in the proliferation of these plants to the extent of their no longer being at risk of extinction. The ability to do the work exists. What is needed is a massive cutting of the swamp of bureaucratic red tape that has resulted from a well-intentioned, idealistic international treaty seriously flawed in the realities of the everyday world!

LEAVES AND CELLS
The plant leaf is a disposable factory. Most factories are a building with machinery installed, raw material supplied, work being done, and a finished product produced. Over time, machines are repaired or replaced, but the basic building endures. However, plant leaf factories are governed by the five basic phases of biology: beginning, growth, maturity, decline, and death! The leaf starts as a bud, then enlarges into being a fully grown leaf. At maturity, it does its work as a factory. The leaf utilizes water and minerals supplied by the roots and through open stomata take in carbon dioxide from the air from the air, while providing for the elimination of oxygen as waste material. With energy supplied by sunlight, the leaf produces the sugars which are transported by a vascular system which feeds all other plant parts. The leaf eventually ages, declines in ability to provide nutrition to the plant system, ultimately dies, is discarded by the plant system, and a whole new leaf factory is produced. Thus, while the leaf is a nutrition producing factory for the plant, it becomes not a disposable machine, but a disposable factory! Plant Cells Some plants have relatively large leaf cells while others have small-to-relatively-minute ones. To understand why this is so, it may be useful to consider a short review of plant evolution. Assume a large pool filled with water in which are balloons filled, for the most part, with water, but each one with a small amount of oil or air. Some balloons are filled to their capacity, but others less so. In the pool, all float, and are relatively weightless, much like small fishes or whales in an ocean. However, when a plug is pulled and the water is drained from the pool all these ―cells‖ are stranded on the land surface. Now the walls of the larger cells are subjected to much more stress than those of the smaller ones, so the walls of the larger cells tend to break. The same may be noted with plant cells. As plant genera progress from tropical to temperate zone environments the climatic selection pressures become progressively greater. Likewise, the same may be said for plants spreading from wet to dry environments, shady to sunlit environments, and so forth. This was graphically illustrated to this worker while investigating the leaf cells of members of the genus Stanhopea, an orchid genus that originated in the New World tropics and-over time-radiated into near-temperate zones. The tropical members of Stanhopea have relatively large leaf cells, but as the genus radiated, the leaf cells became smaller as speciation progressed. Much more might be explored concerning this topic, but for this work it is sufficient to note that as plant cells have relatively rigid walls, any expansion and contraction activity will be correspondingly greater in larger celled leaves, and will necessitate greater cushioning between any expanding members and the rigid cell walls surrounding them, with the reverse being true of smaller-celled leaves. Adaxial and Abaxial Surfaces In many mature leaves, the upper (adaxial) surface (the one exposed to the sunlight) is covered with a waxy coating. This acts as not only a form of sunscreen, but aids in shedding rain so the leaf cells don‘t become overly saturated with water and burst. This upper surface may look like just so many bricks with an occasional wax-secreting gland present (Fig. 1). The adaxial surface may even sprout prickles, thorns or hairs which may aid in deterring large to even tiny predators (Fig. 2). The underside of the leaf (the abaxial side) is a different story. Many of these
169

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

surfaces look like just so many cells rather irregularly arranged, and punctuated with small openings looking much like a pair of bananas joined at their ends. As these ―bananas‖ shrink, an opening into the leaf‘s interior is provided. When they swell, the opening into the interior of the leaf is closed. These specialized cells are known as guard or stomata cells because when they‘re open, the opening looks something like a ―stoma‖ (mouth) into the plant interior (Fig. 3). Likewise, these specialized cells act to ―guard‖ the ―mouth‖ entrance into the plant‘s interior.

Fig. 1. Abaxial epidermal leaf cells, x320, Stanhopea oculata. Pl#290496-5 C6. Xylol strip Abα2. 26 May, 1996. 35mm Photomicrograph transparency 19 July, 1996.

Fig. 2. Abaxial epidermal leaf cells, x80, Stanhopea oculata. Pl#290496-5 C6. Xylol strip Abα-2. 26 May, 1996. 35mm Photomicrograph transparency 19 July, 1996.

Guard and Subsidiary Cells Land plants may have stomata on one or both leaf surfaces. Stomata are common on the abaxial surface; but-when present-are usually more sparse on the adaxial leaf surface. The guard cells are very different from other epidermal cells, and subsidiary cells are also structurally unlike the surrounding epidermal cells, and may be quite distinctive. Certain plant groups have additional cells arranged around their stomatal cells, but in some genera no additional specialized cells are present. These additional specialized cells are referred to as subsidiary or accessory cells. They may act to assist, reinforce, or protect the stomatal cells. Given that plant cells have relatively rigid cellulose cell walls, and that the stomata must expand and contract, subsidiary cells afford a cushioning effect for the adjoining (more rigid) cells from the stomatal expansions and contractions. Subsidiary cells have been noted by this worker as more prevalent in plants with larger leaf cells, and less so in ones with small-to-tiny leaf cells. This evolutionary adaption may have evolved, in part, because the Fig. 3. Stomata guard cells. Drawing, 01 expansion and contraction of the stomata in the smaller February, 2008. celled plants does not generate comparable stresses on the adjoining leaf cells. N. H. Williams (1979) suggests the presence of four subsidiary cells with each stomata may be a derived condition, but outgroup comparison suggests the presence of subsidiary cells as an ancestral condition for the Orchidaceae. The pattern in the Cranichideae is distinctive, and may be characterized by the mesoperigenous development of the subsidiary cells (Williams, 1975).
170

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

Dressler (1993) states the ontogeny of subsidiary cells is considered important in plant clasification, but he instantly qualifies this statement by noting that there are many systems of classifying stomatal types, each with its own abstruse terminology. Clinical research by this worker (1999), utilizing extensive measuring of guard and subsidiary cells, found no statistical support for these cells having taxonomic significance as regards species identification in the genus Stanhopea. Likewise, no similar taxonomic significance was found in other orchids studied, albeit research was less exhaustive with members of Govenia, Laelia, Malaxis, Phalaenopsis, and other genera. Dressler‘s offering on stomata and subsidiary cells is limited to less than a page (three paragraphs), in which he notes the following, ―As is too often the case, we have good information on the pattern and ontogeny of the epidermis in the advanced Epidendroideae, but more study of the primitive groups is needed.‖ (page 23)

Fig. 4. Anisocytic Subsidiary cells. Drawing, 01 February, 2008.

Fig. 5. Paracytic Subsidiary cells. Drawing, 01 February, 2008.

In the course of researching guard and subsidiary cells, and reviewing the classification systems of other workers, this worker found it useful to simplify the classification of subsidiary cell formations into three groups: anisocytic, paracytic, or diacytic. Each of these three groups does essentially the same job and is composed of the same cellular material, but these cells differ from the composition of the guard cells and the other normal epidermal leaf cells. Anisocytic cells (aniso: ―unequal) are unequal in appearance to each other (Fig. 4). An anisocytic cell group may be composed of three or more cells which surround the guard cells, buffering it from the other epidermal leaf cells. Paracytic (para, around, and cytos: cells, not ―parasitic,‖ which has to do with being a parasite!) subsidiary cells are arranged about the long axes of the stomata cells (Fig. 5). The third group of subsidiary cells are called Diacytic (dia, across) because they are arranged at right angles to the stomata cells (Fig. 6). The bewhiskered subsidiary cells shown in Fig. 2 are clearly paracytic ones, and although they vary in size and frequency from one species to another, this type of subsidiary cell formation is a general characteristic of members of the genus Stanhopea; a decidedly tropical to an almost-temperate genus. Compare these cells with the ones from the abaxial surface of Govenia utriculata, a terrestrial orchid species (Fig. 7). The subsidiary cells of Stanhopea oculata specimen in Fig. 1 averaged 1632.02μ, while those of the terrestrial Govenia superba averaged a
171

Fig. 6. Diacytic Subsidiary cells. Drawing, 01 February, 2008.

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

mere 865.51μ. The guard cells of plants with relatively large epidermal leaf cells obviously require much more ―cushioning‖ than do the guard cells of ones of smaller sized epidermal leaf cells.

Fig. 7. Abaxial epidermal leaf cells, x80, Govenia utriculata. Pl#230796-2 C35. Xylol strip Abα-1: 23 July, 1996. 35mm Photomicrograph transparency digitalized: 31 January, 2008.

Less obvious, albeit not surprisingly, as epidermal leaf cells are smaller, the evolutionary need for ―cushioning‖ of subsidiary cells becomes minimal or may vanish altogether. In fact, in the Orchidoideae, Neottieae, and Pogoniinae, recognizable subsidiary cells are consistently lacking (Dressler, 1993), and H. Rasmussen (1981, 1987) indicates that some cells in the Orchioideae may be mesoperigenous in origin, but not recognizable as subsidiary cells at maturity.

RECENT WORK

Fig. 8. Abaxial epidermal leaf cells, S. stellata. Dried leaf, X80 mag, 35mm transp#5, 19Jan08.

FIG. 9 Abaxial epidermal leaf cells, S. stellata. Dried leaf, X320 mag, 35mm transp#7, 19Jan08

172

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

Guard cells were clearly visible at X80 magnification, but no subsidiary cells were evidenced (Fig. 11). The abaxial leaf surface was then examined at X320 magnification (Fig. 12) and again, even more clearly, no subsidiary cells were visible.

FIG. 10 Abaxial epidermal leaf cells, S. romanzoffiana. Dried leaf, X80 mag, 35mm transp#16, 19Jan08

FIG. 11 Abaxial epidermal leaf cells, S. cernua. Live leaf, X80 mag, 35mm transp#23, 19Jan08

Fig. 12. Abaxial epidermal leaf cells, Spiranthes cernua. Live leaf, X320 mag, 35mm transparency #24. 19Jan08.

DISCUSSION, CONCLUSIONS, AND OUTLOOK
No other species of Spiranthes were examined, but from examining adaxial and abaxial specimens of Spiranthes stellata, S. romanzoffiana, and S. cernua, it is inferred that at least at maturity, subsidiary cells are absent in members of this genus. In no case were stomata observed on the adaxial leaf surfaces of the Spiranthes specimens examined in this limited study. The examination of leaf samples at X32 magnification shows the generalized cell pattern for both the adaxial and abaxial surfaces. However, if critical measuring of epidermal leaf cells is to be done, as this worker did extensively with members of the genus Stanhopea, magnification at X80 is recommended. Few specimens were examined in this study, and it may be that taxonomically significant patterns would emerge if a large number of species of several genera were surveyed and the data submitted to an analysis of variance treatment (ANOVA). However, in order to ensure statistical validity, this worker would insist on an absolute minimum of five samples of of each species, with ten randomly collected specimens of each species considered as a more robust sampling. If

173

Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS

this work is done, it is suggested that data from the adaxial leaf cells probably will be found to be more significant from a taxonomic standpoint than the data collected from abaxial leaf surfaces. For general use by field biologists, for example, in a national park environment, it would be useful to have access to certain readily confirmed general identification markers such as the presence or absence of subsidiary cells, or the general appearance of wax glands on the adaxial leaf surface or the appearance and general frequency of stomatal guard cells on adaxial leaf surfaces. If so, these could be additional tools for the recognition of certain species during nonflowering periods. REFERENCES
Dressler, R.L. 1993. Phylogeny and Classification of the Orchid Family. Portland, Oregon: Dioscorides Press. Ferry Townsend, R.J. 1999. Estudio Anatómico Epidermico en Las Hojas Del Genero Stanhopea (Orchidaceae) y Sus Implicaciones Taxonómicas. Monterrey, NL, México: Universidad Autónoma de Nuevo León. 49pp y Anexo I (68pp.), II (27pp.), III (44pp.), y Epilogo (1p.). Rasmussen, H. 1981. The diversity of stomatal development in Orchidaceae subfamily Orchidoideae. Botanical Journal of the Linnean Society 82: 381-93. _______ . 1987. Orchid stomata-structure, differentiation, function, and phylogeny. In Orchid Biology, Reviews and Perspectives, Vol. 4. Ed. J. Arditti. Ithaca: Cornell University Press. Stewart S.L. and L.W. Zettler, 2002. Symbiotic germination of three semi-aquatic rein orchids (Habenaria repens, H. quinqueseta, H. macroceratitis) from Florida. Aquatic Botany: 72: 25-35. Stewart S.L., L.W. Zettler, J. Minso, and P.M. Brown. 2003. Symbiotic germination and reintroduction of Spiranthes brevilabris Lindley, an endangered orchid native to Florida. Selbyana: 24: 64-70. Stewart S.L., M.E. Kane. 2006. Symbiotic seed germination of Habenaria macroceratitis (Orchidaceae), a rare Florida terrestrial orchid. Plant Cell Tissue Organ Culture 86: 159-67. Williams, N.H. 1975. Stomatal Development in Ludisia discolor (Orchidaceae): mesoperigenous subsidiary cells in the mocotyledons. Taxon 24: 281-88. _______. 1979. Subsidiary cells in the Orchidaceae: their general distribution with special reference to development in the Oncidieae. Botanical Journal of the Linnean Society 78: 41-66. Zettler, L.W. 1997a. Orchid fungal symbiosis and its value in conservation. McIlvania. 13: 40-45. _______. 1997b. Terrestrial orchid conservation by symbiotic seed germination: techniques and perspectives. Selbyana 18: 188-94. Robert J. Ferry, PhD, 343 John Wayne Trail, Victoria, Texas 77905 email: rferry@miosjournal.org

174

Empiricist: STAYING HEALTHY Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

STAYING HEALTHY
The Slow Empiricist
As people in the United States are aging I am inspired to write about how having an interest like native orchids can be of great benefit to your staying healthy and connected to the world around you. Just this week I read that if you want to live to be 100 years old you need to be active and involved as you age. See the March 10, 2008 copy of the Parade magazine about some of the things that keep you engaged and healthy.

Stan Folsom with Spiranthes ochroleuca in Nova Scotia

If you have a passion such as orchid hunting and photographing you are already on the road to longevity in my estimation. Some of the benefits of this enterprise are the actual physical activity of getting out into the field and hiking and searching for the plants. This requires physical exercise and mental prowess as you
175

Empiricist: STAYING HEALTHY Stewart & Richardson: FLORIDA PANTHER NWR ORCHID FLORA

do your explorations. You also will probably find that you are not alone in your pursuits and you may end up with some enduring friendships along your way. Even in the months that you can't get out into the field because of the winter prohibitions you can engage your mind in reading about the plants or in attending programs offered by the many orchid societies that abound in the United states and Canada, or, for that matter in anywhere in the world for orchid enthusiasts are everywhere. If you are fortunate enough to afford travel to warmer climates you might plan and take a trip to explore for the plants in a different locale than your own backyard. All this activity is good for your brain and your body to experience. There are some constraints that you need to be aware of before you start out to conquer the orchid world. If you have some disability that might affect you negatively you should make serious plans to circumvent any adverse affects your problem might throw at you. I just learned that I have borderline diabetes so now I must take that into consideration when I am out in the field. I don't expect it to be a major impediment to my enjoying a good tramp in the woods or across the meadows in search of orchids but I will now make sure I carry snacks to give me proper energy and drinks to keep me hydrated. I will also take along my cell phone and hope I have cells if there is an actual emergency. My best protection is to make sure that I am never alone on my explorations. I have heard too many horrendous escapades that have occurred from people who think they are invincible and set out alone to do their orchid explorations. They get lost, they fall and injure themselves, and they encounter less than friendly wild things that jeopardize their survival. Part of growing old gracefully is to recognize your limitations and not try to be Superman or woman. Now if you get to the point where being in the field is really just too hard for you to do any more, may I suggest that you continue to be active in whatever way that you can. Maybe you can start compiling your memories and thoughts about what you experienced in your better days. You could augment your written words with photos from your experiences and you are on the way to creating a viable program for your friends and acquaintances to enjoy. You could share them with people by presenting a program for your local orchid society or you could offer your memoirs to the North American Native Orchid Journal so that our readers could enjoy your experiences, too. Your memories could expand the knowledge we have about native orchids and enrich a whole new generation of orchid enthusiasts. What a legacy!

Your Slow Empiricist

176

Brown: HYBRIDS IN THE GENUS CALOPOGON

HYBRIDS IN THE GENUS CALOPOGON
Paul Martin Brown One of the showiest genera of orchids to be found in North America, Calopogon, the grass-pinks, is comprised of five species. Calopogon tuberosus var. tuberosus is one of the most wideranging species of orchid found in eastern North America, C. oklahomensis while currently restricted to remnant prairies in the Midwest, is known from historical collections ranging over a much wider area. The remaining three species, C. multiflorus, C. barbatus, and C. pallidus are primarily southeastern coastal plain species. A variety of C. tuberosus, var. simpsonii, is restricted to southern Florida marls and also found in nearby Bahamas and Cuba. Hybrids among the four of the five species that often grow sympatrically are not all that unusual, although often difficult to detect. Calopogon oklahomensis, itself a species of hybrid origin, grows and flowers at a time in range and habitats when no other Calopogon is present. The works of Goldman (2000), Goldman and Orzell (2000); Goldman et al. (2004); Trapnell (1994); Trapnell et al. (2004). have paved the way for this assessment of the hybrids. Goethe State Forest in Levy County Florida presents the four southeastern species growing sympatrically and most years they have an overlap of flowering time. It is here that the six hybrids can be more easily detected (Fig. 1). Depending on the combination of parents some hybrids are not difficult to determine while in other cases it is more problematic. It is not that unusual to find two species growing together, rarely three, but all four in such a concentrated area has both encouraged and allowed for detail examination of the potential hybrids. The following are proposed as names for the six hybrid combinations: Calopogon xfloridensis P.M. Brown nothosp. nov Planta inter Calopogon multiflorus et Calopogon pallidus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon multiflorus and Calopogon pallidus TYPE: U.S.A. Florida: Highlands Co.: Avon Park AFR. 17 April 1997. Goldman & Orzell 1291. (Holotype TEX), photograph Fig. 5b
PARATYPES:U.S.A. Florida: Hillsborough Co. near Seaside, 8 May 1899. Barnhardt 2763 (NY); Duval Co., Apr. Curtiss 2801 (MO); Martin Co.: Jonathan Dickinson State Park, Jan. 1994 Goldman & Goldman 1442 (TEX, photo); Pinellas Co., Tarpon Springs, Mar. 1921, Beckwith 605A (ROCH)

Etymology: the epithet floridensis is chosen because of the first observation in the state of Florida Common name: Florida hybrid grass-pink Plants of Calopogon xfloridensis are well-documented by Goldman & Orzell (2000). They cite five locations for this hybrid and additional plants have been seen by this author in three northern Florida and southern Georgia locations. Calopogon xobscurus P.M. Brown nothosp. nov Planta inter Calopogon barbatus et Calopogon multiflorus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon barbatus and Calopogon multiflorus

177

Brown: HYBRIDS IN THE GENUS CALOPOGON

Type: U.S.A. Florida: Wakulla Co., Ochlockonee River State Park. 10 April 2005 P.M. Brown 0514 (Holotype: Ochlockonee River State Park Herbarium) photograph Fig. 4b Etymology: the epithet obscurus is chosen because the nature of the hybrid as it ‗hides‘ or is obscure among the parents Common Name: hidden hybrid grass-pink Although less frequently sharing habitat C. multiflorus is abundant in seasonally burned pine flatwoods in Goethe State Forest and it is not difficult to find populations of C. barbatus nearby and occasional plants of Calopogon xobscurus clearly intermediate between the two putative parents Additional plants of this hybrid have been seen at Ochlockonee River State Park, where the holotype was collected and in southern Georgia. Calopogon xvulgaris P.M. Brown nothosp. nov Planta inter Calopogon pallidus et Calopogon tuberosus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon pallidus and Calopogon tuberosus Type: U.S.A. Florida: Levy Co., CR 336, Goethe State Forest 15 May 2005 P.M. Brown 05-27 (Holotype: Goethe State Forest Herbarium) photograph Fig. 2b Etymology: the epithet vu;garis is chosen because the hybrid is relatively common where both parents occur Common name: common hybrid grass-pink The most abundant species on the wet roadsides is Calopogon tuberosus, while scattered plants of C. pallidus occur within these populations. It is here that the hybrid Calopogon xvulgaris frequently occurs and dramatically blends the morphology of both parents. Additional plants of this hybrid have been seen frequently throughout central and northern Florida. Calopogon xsimulans P.M. Brown nothosp. nov Planta inter Calopogon barbatus et Calopogon tuberosus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon barbatus and Calopogon tuberosus Type: U.S.A. Florida: Levy Co., CR 336, Goethe State Forest 29 April 2005 P.M. Brown 05-20 (Holotype: Goethe State Forest Herbarium) photograph: Fig. 3b Etymology: the epithet simulans is chosen because the hybrids simulate the species C. oklahomensis Common name: False Oklahoma hybrid grass-pink Perhaps the most surprising hybrid is know from only a handful of plants at Goethe and this is the combination of C. barbatus and C. tuberosus, Calopogon xsimulans, and appearing much like C. oklahomensis. In fact, the first time these were observed several of us suspected the latter species was present there. Then we started looking at the various Calopogon hybrids in the area and realized we had a contemporary hybrid of the same parentage as C. oklahomensis! Calopogon xfowleri P.M. Brown nothosp. nov Planta inter Calopogon barbatus et Calopogon pallidus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon barbatus and Calopogon pallidus Type: U.S.A. Florida: Levy Co., CR 336, Goethe State Forest 30 April 2005 P.M. Brown 05-23 (Holotype: Goethe State Forest Herbarium) photograph: Fig. 5b Etymology: named for Jim Fowler who brought this hybrid to my attention in 2007 from the Green Swamp in southeastern North Carolina Common name: Fowler‘s hybrid grass-pink Calopogon xfowleri is the fifth combination and produces a very beautiful plant that has been seen several places in Florida, Georgia, and North Carolina.
178

Brown: HYBRIDS IN THE GENUS CALOPOGON

Calopogon xgoethensis P.M. Brown nothosp. nov

Planta inter Calopogon multiflorus et Calopogon tuberosus intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Calopogon multiflorus and Calopogon tuberosus Type: U.S.A. Florida: Levy Co., CR 336, Goethe State Forest 30 April 2005 P.M. Brown 05-24 (Holotype: Goethe State Forest Herbarium) photograph: Fig. 7b Etymology: named for Goethe State Forest where the hybrids were first detected The sixth hybrid combination at Goethe, Calopogon xgoethensis, is between the two most abundant parents to be found in the forest region, Calopogon tuberosus and C. multiflorus, but has apparently produced very few hybrid plants. This may be because the bulk of C. multiflorus has finished flowering by the time C. tuberosus are at their peak, although there are always later and early plants of the respective species in flower at the same time. The plants grow within sight of each other in differing habitats. This same cross has been registered as the horticultural hybrid Calopogon Fluffy by Carson Whitlow in 2003.

C. tuberosus

C. pallidus

C. xvulgaris

C. xgoethensis

C. xsimulans

C. xfloridensis

C. xfowleri

C. xobscurus C. barbatus
Figure 1. Calopogon hybrids and parentage
179

C. multiflorus

Brown: HYBRIDS IN THE GENUS CALOPOGON

That these six hybrids have not been described before may be a bit surprising, but it takes a second or even a third look sometimes to detect such plants, especially when the parents are relatively abundant. All six hybrids have been found in an area of not more than 500 feet along the same roadway or within 500 feet of the wood edge.

REFERENCES: Goldman, D.H. 2000. Systematics of Calopogon R. Br. and the tribe Arethusae (Orchidaceae). Ph.D. Dissertation. University of Texas. Goldman, D.H. and L.S. Orzell. 2000. Morphological, geographical, and ecological re-evaluation of Calopogon multiflorus (Orchidaceae). Lindleyana 15:237-51. Goldman, D.H., C. van den Berg, and M.P. Griffith. 2004. Morphometric circumscription of species and infraspecific taxa in Calopogon R.Br. (Orchidaceae) Plant Systematics and Evolution 37-60. Trapnell, D.W. 1994. Systematic review of Calopogon (Orchidaceae). Master‘s thesis. University of Georgia. Trapnell, D.W., J.L. Hamrick, and D.E. Giannasi. 2004. Genetic variation and species boundaries in Calopogon (Orchidaceae). Systematic Botany 29: 308–15. ACKNOWLEDGEMENTS: The author wishes to thank Scott Stewart for helpful comments and field assistance and Wally Wilder for observations in the field.

Fig. 2 C. pallidus

C. xvulgaris

C. tuberosus

Fig. 3 C. tuberosus

C. xsimulans

C. barbatus

180

Brown: HYBRIDS IN THE GENUS CALOPOGON

Fig. 4 C. barbatus

C. xobscurus

C. multiflorus

Fig. 5 C. multiflorus

C. xfloridensis

C. pallidus

Fig. 6 C. barbatus

C. xfowleri

C. pallidus

Fig. 7 C. multiflorus

C. xgoethensis
181

C.. tuberosus

BOOK REVIEWS & PUBLICATION ANNOUNCEMENTS NEW TAXA AND COMBINATIONS IN THIS ISSUE Calopogon xfloridensis P.M. Brown nothosp. nov. p. 176 Calopogon xobscurus P.M. Brown nothosp. nov. p. 176 Calopogon xvulgaris P.M. Brown nothosp. nov. p. 176 Calopogon xsimulans P.M. Brown nothosp. nov. p. 176 Calopogon xfowleri P.M. Brown nothosp. nov. p. 176 Calopogon xgoethensis P.M. Brown nothosp. nov. p. 176 Galearis spectabilis (L.) Raf. forma lilacina (Ames) P.M. Brown stat. & comb. nov. p. 138

182

BOOK REVIEWS & PUBLICATION ANNOUNCEMENTS

Available from your favorite bookseller, University of Iowa Press, or the authors at naorchid@aol.com These are the first two in a projected series of 12 guides.
183

BOOK REVIEWS & PUBLICATION ANNOUNCEMENTS

NOW AVAILABLE!
Ideal for the casual or dedicated orchid enthusiast "Clear and concise text, easy-touse keys, breathtaking photography, exquisite line drawings, and easily tucked into your backpack! What more could one ask of this nifty field guide?"--Helen Jeude, Botanical Research Institute of Texas "An outstanding addition to previous similar works by the same author-artist team. However, this work and the others are much more than mere 'field guides!' They clarify and update information regarding the orchid flora of the United States, and for the orchid hobbyist-hunter, the 'where-tolook-for-orchids' portion alone may be worth more than the price of the book."--Dr. Robert J. Ferry, McAllen (Texas) International Orchid Society Though many may not realize it, Texas is home to some of the rarest orchid species in the world. From the Big Thicket to the Big Bend, from the Panhandle to the coastal plains, Field Guide to the Wild Orchids of Texas is the first native orchid field guide to cover each of the Lone Star State's eco-regions. 60 species and varieties – 4 new to Texas 38 color and growth forms -- 4 recently published and new to Texas; 8 hybrids; details on 17 additional species that grow adjacent to the borders of Texas and may eventually be found there University Press of Florida - $29.95 - ISBN13:978-0-8130-3159-0 316 pages; 288 color photographs; 6 color watercolors; 91 line drawings; 67 full color maps

Signed and inscribed copies from the authors at naorchid@aol.com The perfect compliment and companion to the Liggio‘s Wild Orchids of Texas (1999).
184