June 2008 North American Native Orchid Journal

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………….
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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.
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NORTH AMERICAN
NATIVE ORCHID JOURNAL
Volume 14 (2) 2008
CONTENTS
NOTES FROM THE EDITOR

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NEWS OF INTEREST:
SPIRANTHES DILUVIALIS NEW TO CANADA
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ORCHID FLORA OF THE
FLORIDA PANTHER NATIONAL WILDLIFE REFUGE
Scott L. Stewart & Larry W. Richardson
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SYNONYMS AND MISAPPLIED NAMES
OF WILD ORCHIDS FOUND IN
NORTH AMERICA NORTH OF MEXICO
105
A NEW COMBINATION
138
MY FAVORITE THINGS
Connie Bransilver
139
THE SPIRANTHES FORMERLY KNOWN AS PARKSII
Lucy A. Dueck
143
PLATANTHERA PALLIDA, PALE FRINGED ORCHIS
Robert T. McGrath
150
COMING IN JULY……
157
PLATANTHERA PALLIDA - FIFTEEN YEARS OF COMPARISONS
Paul Martin Brown
158
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GUARD CELLS IN SPIRANTHES
Robert J. Ferry
167
STAYING HEALTHY
The Slow Empiricist
174
HYBRIDS IN THE GENUS CALOPOGON
Paul Martin Brown
176
NEW TAXA AND COMBINATIONS IN THIS ISSUE
181
BOOK REVIEWS & PUBLICATION ANNOUNCEMENTS
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
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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
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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.
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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‖
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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 twenty-
seven 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
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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.
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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. A wet prairie on the FPNWR with spider
lilies (Hymenocallus palmeri).
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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.
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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.
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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 pink-
purple 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.
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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.
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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.
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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.
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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.
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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.
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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.
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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.
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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.
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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 refuge-
wide orchid conservation program.
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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.
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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.
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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.
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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.
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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.
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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
Scientific name: Harrisella porrecta (Reichenbach f.) Fawcett & Rendle

Synonyms: Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten
Harrisella filiformis (Swartz) Cogniaux – misapplied
Common name: leafless harrisella
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.
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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
Scientific name: Oeceoclades maculata (Lindley) Lindley

Common name: spotted African orchis
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.
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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
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
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
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
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.
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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.
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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; 10-
30 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 yellow-
throated 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.
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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.
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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.
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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
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. Larry W. Richardson

PhytoTechnology Laboratories, LLC Florida Panther National Wildlife
Lenexa, KS Refuge
scott@phytotechlab.com Naples, FL
larry_richardson@fws.gov
104
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: Orchisrotundifolia 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
Aplectrum hyemale (Mühlenberg in Willdenow) Nuttall
BASIONYM: Cymbidium hyemale Mühlenberg in Willdenow
SYNONYM: Aplectrum spicatum Britton, Sterns & Poggenberg
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forma pallidum (House) Barnhart

BASIONYM: Aplectrum spicatum Britton, Sterns & Poggenberg forma pallidum House
Basiphyllaea corallicola (Small) Ames
BASIONYM: Carteria corallicola Small
Beloglottis costaricensis (Reichenbach f.) Schlechter
BASIONYM: Spiranthes costaricensis Reichenbach f.
Bletia purpurea (Lamark) de Candolle
BASIONYM: Limodorum purpureum Lamark
forma alba (Ariza-Julia & J. Jiménez Almonte) P.M. Brown
BASIONYM: Bletia purpurea (Lamark) de Candolle var. alba Ariza-Julia & J. Jiménez Almonte
Brassia caudata (Linnaeus) Lindley
BASIONYM: Epidendrum caudatum Linnaeus
Bulbophyllum pachyrhachis (A. Richard) Grisebach
BASIONYM: Pleurothallis pachyrhachis A. Richard
Calopogon barbatus (Walter) Ames
BASIONYM: Ophrys barbata Walter
SYNONYMS: Calopogon graminifolius Elliot in Weatherby & Griscom
Calopogon parviflorus Lindley
Calopogon pulchellus (Salisbury) R. Brown var. graminifolius Elliot
Limodorum parviflorum (Lindley) Nash
Calopogon multiflorus Lindley
SYNONYMS: Calopogon barbatus (Walter) Ames var. multiflorus (Lindley) Correll
Limodorum multiflorum (Lindley) Mohr
Limodorum pinetorum Small
Calopogon pallidus Chapman
SYNONYM: Limodorum pallidum (Chapman) Mohr
Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. tuberosus
BASIONYM: Cathea tuberosa (Linnaeus) Salisbury
SYNONYMS: Calopogon pulchellus (Salisbury) R. Brown
Calopogon pulchellus (Salisbury) R. Brown var. latifolius (St. John) Fernald
Limodorum pulchellum Salisbury
Limodorum tuberosum Linnaeus
Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var. simpsonii (Small)
Magrath
BASIONYM: Limodorum simpsonii Small
SYNONYM: Calopogon pulchellus (Salisbury) R. Brown var. simpsonii (Small) Ames
Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown) Luer
BASIONYM: Calypso americana R. Brown in Aiton
SYNONYM: Cytherea bulbosa (Linnaeus) House p.p.
Calypso bulbosa (Linnaeus) Oakes var. occidentalis (Holtzinger) Boivin
BASIONYM: Calypso bulbosa (Linnaeus) Oakes forma occidentalis Holzinger
SYNONYMS: Calypso bulbosa Britton subsp. occidentalis (Holzinger) Calder & Roy L. Taylor
Calypso occidentalis (Holzinger) A. Heller
Cytherea bulbosa House var. occidentalis (Holzinger) Cockerell
Cytherea occidentalis (Holzinger) A. Heller
Cephalanthera austiniae (A. Gray) Heller
BASIONYM: Chloraea austinae A. Gray
SYNONYMS: Cephalanthera oregana Reichenbach
Eburophyton austinae (A. Gray) Heller
Epipactis austinae (A. Gray) Wettstein
Serapias austinae (A. Gray) A.A. Eaton
Cleistes divaricata (Linnaeus) Ames
BASIONYM: Arethusa divaricata Linnaeus
SYNONYM: Pogonia divaricata (Linnaeus) R. Brown
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Cleistes bifaria (Fernald) Catling & Gregg

BASIONYM: Cleistes divaricata (Linnaeus) Ames var. bifaria Fernald
SYNONYM: Pogonia bifaria (Fernald) P.M. Brown & Wunderlin
Coeloglossum viride (Linnaeus) Hartman var. viride
BASIONYM: Satyrium viride Linnaeus
SYNONYMS: Coeloglossum viride (Linnaeus) Hartman var. islandicum (Lindley) Schulze
Dactylorhiza viridis (Linnaeus) R.M. Bateman, A. Pridgeon & M.W. Chase
Habenaria viridis (Linnaeus) R. Brown in Aiton f.
Coeloglossum viride (Linnaeus) Hartman var. virescens (Mühlenberg) Luer
BASIONYM: Orchis bracteata Mühlenberg in Willdenow
SYNONYMS: Coeloglossum bracteatum (Mühlenberg in Willdenow) 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
Corallorhiza mertensiana Bongard
SYNONYMS: 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
var. maculata forma flavida (Peck) Farwell
BASIONYM: Corallorhiza multiflora Nuttall var. flava Peck
Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley) Ames
BASIONYM: Corallorhiza multiflora Nuttall var. occidentalis Lindley
SYNONYM: Corallorhiza maculata (Rafinesque) Rafinesque subsp.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
Corallorhiza maculata var. mexicana (Lindley) J.V. Freudenstein
BASIONYM: Corallorhiza mexicana Lindley
Corallorhiza odontorhiza (Willdenow) Poiret var. odontorhiza
BASIONYM: Cymbidium odontorhizum Willdenow
SYNONYM: Corallorhiza micrantha A. Chapman
Corallorhiza odontorhiza (Willdenow) Poiret var. pringlei (Greenman) Freudenstein
BASIONYM: Corallorhiza pringlei Greenman
Corallorhiza striata Lindley var. striata
SYNONYMS: Corallorhiza macraei A. Gray
Neottia striata (Lindley) Kuntze
Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams
BASIONYM: Corallorhiza vreelandii Rydberg
SYNONYMS: Corallorhiza bigelovii S. Watson
Corallorhiza ochroleuca Rydberg
forma flavida (Todsen & Todsen) P.M. Brown
BASIONYM: Corallorhiza striata Lindley var. flavida T.A. Todsen & Todsen
Corallorhiza trifida Chatelain
SYNONYM: Corallorhiza corallorhiza (Linnaeus) Karsten
forma verna (Nuttall) P.M. Brown
BASIONYM: Corallorhiza verna Nuttall
SYNONYM: Corallorhiza trifida Chatelain var. verna (Nuttall) Fernald
Corallorhiza wisteriana Conrad
SYNONYMS: Corallorhiza odontorhiza sensu Chapman not Nuttall
107
Corallorhiza unguiculata Rafinesque

forma albolabia P.M. Brown
SYNONYM: Corallorhiza wisteriana Conrad forma toleri S. Bentley
Cyclopogon cranichoides (Grisebach) Schlechter
BASIONYM: Pelexia cranichoides Grisebach
SYNONYMS: Beadlea cranichoides (Grisebach) Small
Spiranthes cranichoides (Grisebach) Cogniaux
Cyclopogon elatus (Swartz) Schlechter
BASIONYM: Satryrium elatum Swartz
SYNONYMS: Beadlea elata (Swartz) Small
Spiranthes elata (Swartz) L.C. Richard
Cypripedium acaule Aiton
SYNONYM: Fissipes acaulis (Aiton) Small
Cypripedium arietinum R. Brown
SYNONYM: Criosanthes arietina (Aiton f.) House
Cypripedium kentuckiense C.F. Reed
Cypripedium daultonii V. Soukup nom. nud.
Cypripedium fasciculatum Kellogg in S. Watson
SYNONYM: Cypripedium knightae A. Nelson
Cypripedium montanum Douglas in Lindley
SYNONYM: Cypripedium occidentale Watson
Cypripedium parviflorum Salisbury var. parviflorum
SYNONYM: Cypripedium calceolus Linnaeus var. parviflorum (Salisbury) Fernald p.p.
Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak
BASIONYM: Cypripedium pubescens Willdenow var. makasin Farwell
SYNONYM: Cypripedium calceolus Linnaeus var. parviflorum (Salisbury) Fernald p.p.
Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight
BASIONYM: Cypripedium pubescens Willdenow
SYNONYMS: Cypripedium calceolus Linnaeus var. 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
Cypripedium passerinum Richardson forma minganense (Victorin) P.M. Brown
BASIONYM: Cypripedium passerinum Richardson var. minganense Victorin
Cypripedium reginae Walter
SYNONYM: Cypripedium spectabile Salisbury
Cypripedium yatabeanum Makino
SYNONYMS: Cypripedium guttatum subsp. yatabeanum (Makino) Hultén
Cypripedium guttatum var. yatabeanum (Makino) Pfitzer
Cyrtopodium macrobulbon (La Llave & Lexarza) G.A. Romero & Carnevali
BASIONYM: Epidendrum macrobulbon La Llave & Lexarza
Cyrtopodium polyphyllum (Vellozo) Pabst in F. Barrios
BASIONYM: Epidendrum polyphyllum Vellozo
MISAPPLIED NAME: Cyrtopodium andersonii (Lambert in Andrews) R. Brown in Aiton
Cyrtopodium punctatum (Linnaeus) Lindley
BASIONYM: Epidendrum punctatum Linnaeus
Dactylorhiza aristata (Fischer in Lindley) Soó
BASIONYM: Orchis aristata Fischer in Lindley
Dactylorhiza aristata (Fischer in Lindley) Soó var. kodiakensis Luer forma perbracteata
(Lepage) P.M. Brown
BASIONYM: Orchis aristata forma perbracteata Lepage
108
SYNONYM: Dactylorhiza aristata (Fischer ex Lindley) Soó forma perbracteata (Lepage) Catling
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
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
Dactylorhiza majalis (Reichenbach f.) Summerhayes var. praetermissa (Druce) D.M.
Moore & Soó *?
BASIONYM: Orchis praetermissa Druce
SYNONYM: Dactylorhiza praetermissa (Druce) Soóvar. praetermissa
forma albiflora (Druce) P.M. Brown
BASIONYM: Orchis praetermissa Druce var. albiflora Druce
Deiregyne confusa Garay
SYNONYMS: 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
Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten
BASIONYM: Aeranthes porrectus Reichenbach f.
SYNONYMS: Campylocentrum porrectum Reichenbach f.
Harrisella porrecta (Reichenbach f.) Fawcett & Rendle
Dendrophylax lindenii (Lindley) Bentham in Rolfe
BASIONYM: Angraecum lindenii Lindley
SYNONYMS: Polyradicion lindenii (Lindley) Garay
Polyrrhiza lindenii (Lindley) Cogniaux
Dichromanthus cinnabarinus (La Llave & Lexarza) Garay
BASIONYM: Neottia cinnabarina La Llave & Lexarza
SYNONYMS: Spiranthes cinnabarina (La Llave & Lexarza) Hemsley
Stenorrhynchos cinnabarina (La Llave & Lexarza) Lindley
Dichromanthus michuacanus (La Llave & Lexarza) Salazar & Soto-Arenas
BASIONYM: Neottia michuacana La Llave & Lexarza
SYNONYMS: Spiranthes michuacana (La Llave & Lexarza) Hemsley
Stenorrhynchos michuacanum (La Llave & Lexarza) Lindley
Eltroplectris calcarata (Swartz) Garay & Sweet
BASIONYM: Neottia calcarata Swartz
SYNONYMS: Centrogenium setaceum (Lindley) Schlechter
Pelexia setacea Lindley
Encyclia rufa (Lindley) Britton & Millspaugh
BASIONYM: Epidendrum rufum Lindley
SYNONYM: Encyclia bahamensis (Grisebach) Britton & Millspaugh
MISAPPLIED NAME: Epidendrum bahamense Grisebach
Encyclia tampensis (Lindley) Small
BASIONYM: Epidendrum tampense Lindley
Epidendrum acuñae Dressler
MISAPPLIED NAME: Epidendrum blancheanum Urban
Epidendrum amphistomum A. Richard
MISAPPLIED NAME: Epidendrum anceps Jacquin
Epidendrum magnoliae Mühlenberg var. magnoliae
109
SYNONYM: Epidendrum conopseum R. Brown

Epidendrum magnoliae Mühlenberg var. mexicanum (L.O. Williams) P.M. Brown
BASIONYM: Epidendrum conopseum R. Brown var. mexicanum L.O. Williams
Epidendrum floridense Hágsater
MISAPPLIED NAMES: Epidendrum difforme Jacquin
Epidendrum umbellatum Swartz
Neolehmannia difformis (Jacquin) Pabst
Epipactis atrorubens (Hoffman) Besser*
BASIONYM: Serapias latifolia Hudson (rank unspecified) atrorubens Hoffman
Epipactis gigantea Douglas in Hooker
SYNONYMS: Amesia gigantea (Douglas) A. Nelson & Macbride
Helleborine gigantea (Douglas) Druce
Peramium giganteum (Douglas) Salisbury
Epipactis helleborine (Linnaeus) Cranz*
BASIONYM: Serapias helleborine Linnaeus
SYNONYMS: Amesia latifolia (Linnaeus) A. Nelson & 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
Eulophia alta (Linnaeus) Fawcett & Rendle
BASIONYM: Limodorum altum Linnaeus
Galeandra bicarinata G.A. Romero & P.M. Brown
MISAPPLIED NAME: Galeandra beyrichii Reichenbach f.
Galearis spectabilis (Linnaeus) Rafinesque
BASIONYM: Orchis spectabilis Linnaeus
SYNONYM: Galeorchis spectabilis (Linnaeus) 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 oblongifolia Rafinesque
SYNONYMS: 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
BASIONYM: Neottia pubescens Willdenow
SYNONYM: Peramium pubescens (Willdenow) MacMillan
Goodyera repens (Linnaeus) R. Brown
BASIONYM: Satyrium repens Linnaeus
SYNONYM: Goodyera pubescens (Willdenow) R. Brown var. repens (R. Brown) Alphonse Wood
forma ophioides (Fernald) P.M. Brown
BASIONYM: Goodyera repens (Linnaeus) R. Brown var. ophioides Fernald
SYNONYM: Peramium ophioides (Fernald) Rydberg
Goodyera tesselata Loddiges
SYNONYM: Peramium tesselatum (Loddiges) A. Heller
110
Govenia floridana P.M. Brown

MISAPPLIED NAME: Govenia utriculata (Swartz) Lindley
Gymnadenia conopsea (Linnaeus) R. Brown*
BASIONYM: Orchis conopsea Linnaeus
Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata
BASIONYM: Orchis clavellata Michaux
SYNONYMS: Habenaria clavellata (Michaux) 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 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
Gymnadeniopsis integra (Nuttall) Rydberg
BASIONYM: Orchis integra Nuttall
SYNONYMS: Habenaria integra (Nuttall) Sprengel
Platanthera integra (Nuttall) Luer
Gymnadeniopsis nivea (Nuttall) Rydberg
BASIONYM: Orchis nivea Nuttall
SYNONYMS: Habenaria nivea (Nuttall) Sprengel
Platanthera nivea (Nuttall) Luer
Habenaria macroceratitis Willdenow
SYNONYM: Habenaria quinqueseta var. macroceratitis (Willdenow) Luer
Habenaria odontopetala Reichenbach f.
SYNONYMS: Habenaria garberi Porter
Habenaria strictissima Reichenbach f. var. odontopetala (Reichenbach f.) L.O. Williams
Habenella odontopetala (Reichenbach f.) Small
MISAPPLIED NAME: Habenaria floribunda Lindley
Habenaria quinqueseta (Michaux) Eaton
BASIONYM: Orchis quinqueseta Michaux
SYNONYMS: Habenaria michauxii Nuttall
Habenaria simpsonii Small
Platanthera michauxii (Nuttall) Wood
Habenaria repens Nuttall
SYNONYMS: Habenaria nuttallii Small
Platanthera repens (Nuttall) Wood
Harrisella porrecta (Reichenbach f.) Fawcett & Rendle
BASIONYM: Campylocentrum porrectum Reichenbach f.
SYNONYM: Dendrophylax porrectus (Reichenbach f.) Carlsward & Whitten
MISAPPLIED NAME: Harrisella filiformis (Swartz) Cogniaux
Hexalectris grandiflora (A. Richard & Galeotti) L.O. Williams
BASIONYM: Corallorhiza grandiflora A. Richard & Galeotti
SYNONYM: Hexalectris mexicana Greenman
Hexalectris spicata (Walter) Barnhardt
BASIONYM: Arethusa spicata Walter
SYNONYM: Hexalectris aphylla (Nuttall) Rafinesque
Hexalectris spicata var. arizonica (S. Watson) Catling & Engel
BASIONYM: Corallorhiza arizonica S. Watson
Ionopsis utricularioides (Swartz) Lindley
BASIONYM: Epidendrum utricularioides Swartz
Isotria medeoloides (Pursh) Rafinesque
111
BASIONYM: Arethusa medeoloides Pursh

SYNONYMS: Isotria affinis (Austin in A. Gray) Rydberg
Pogonia affinis Austin in A. Gray
Isotria verticillata (Mühlenberg in Willdenow) Rafinesque
BASIONYM: Arethusa verticillata Mühlenberg in Willdenow
SYNONYM: Pogonia verticillata (Mühlenberg in Willdenow) Nuttall
Lepanthopsis melanantha (Reichenbach f.) Ames
BASIONYM: Pleurothallis melanantha Reichenbach f.
SYNONYM: Lepanthes harrissii Fawcett & Rendell
Liparis elata Lindley
MISAPPLIED NAME: Liparis nervosa (Thunberg) Lindley
Listera auriculata Wiegand
SYNONYM: Neottia auriculata (Wiegand) Szlachetko
forma trifolia (Lepage) Lepage
BASIONYM: Listera borealis Morong forma trifolia Lepage
Listera australis Lindley
SYNONYM: Neottia australis (Lindley) Szlachetko
Listera banksiana Lindley
SYNONYMS: Listera caurina Piper
Listera retusa Suksdorf
Neottia caurina (Piper) Szlachetko
Listera borealis Morong
SYNONYM: Neottia borealis (Morong) Szlachetko
Listera cordata (Linnaeus) R. Brown var. cordata
BASIONYM: Ophrys cordata Linnaeus
SYNONYM: Neottia cordata (Linnaeus) Richard
Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hultén
BASIONYM: Listera nephrophylla Rydberg
SYNONYMS: Listera cordata (Linnaeus) R. Brown subsp. nephrophylla (Rydberg) A. & D. Löve
Neottia nephrophylla (Rydberg) Szlachetko
Listera ovata (Linnaeus) R. Brown*
BASIONYM: Ophrys ovata Linnaeus
SYNONYM: Neottia ovata (Linnaeus) Bluff & Fingerhut
Listera smallii Wiegand
SYNONYMS: Listera reniformis Small
Neottia smallii (Wiegand) Szlachetko
Malaxis abieticola Salazar & Soto Arenas
MISAPPLIED NAME: Malaxis tenuis (S. Watson) Ames
Malaxis bayardii Fernald
Malaxis unifolia var. bayardii nom. nud.
Malaxis brachypoda (Gray) Fernald
BASIONYM: Microstylis brachypoda A. Gray
SYNONYM: Malaxis monophyllos (Linnaeus) Swartz var. brachypoda (A. Gray) Morris & Eames
MISAPPLIED NAME: Microstylis monophyllos (Linnaeus) Lindley
forma bifolia (Mousley) Fernald
BASIONYM: Malaxis monophyllos (Linnaeus) Swartz forma bifolia Mousley
Malaxis corymbosa (S. Watson) Kuntze
BASIONYM: Microstylis corymbosa S. Watson
SYNONYM: Achroanthes corymbosa Greene
Malaxis diphyllos Chamisso
SYNONYM: Malaxis monophyllos (Linnaeus) Swartz var. diphyllos (Chamisso) Luer
MISAPPLIED NAMES: Malaxis monophyllos (Linnaeus) Swartz
Microstylis monophyllos (Linnaeus) Lindley
Malaxis paludosa Swartz
112
SYNONYM: Hammarbya paludosa Kuntze

Malaxis porphyrea (Ridley) Kuntze
BASIONYM: Microstylis porphyrea Ridley
MISAPPLIED NAMES: Malaxis ehrenbergii (Reichenbach f.) Kuntze
Malaxis wendtii Salazar
Malaxis spicata Swartz
SYNONYMS: Achroanthes floridana (Chapman) Greene
Malaxis floridana (Chapman) O. Kuntze
Microstylis floridana Chapman
Microstylis spicata (Swartz) Lindley
Malaxis soulei L.O. Williams
SYNONYMS: Achroanthes montana Greene
Malaxis macrostachys (Lexarza) Kuntze
Malaxis montana (Englemann) Kuntze
Microstylis macrostachys Lexarza
Microstylis montana Englemann
Malaxis unifolia Michaux
SYNONYMS: Achroanthes monophylla (Lindley) Greene
Microstylis unifolia (Michaux) Britton, Sterns & Poggenberg
Malaxis wendtii Salazar
SYNONYM: Tamayorkis wendtii (Salazar) R. González & Szlachetko
Maxillaria parviflora (Poeppig & Endlicher) Garay
BASIONYM: Scaphyglottis parviflora Poeppig & Endlicher
SYNONYM: Maxillaria conferta (Grisebach) Schweinfurth in Leon
Mesadenus lucayanus (Britton) Schlechter
BASIONYM: Ibidium lucayanum Britton
MISAPPLIED NAME: Mesadenus polyanthus (Reichenbach f.) Schlechter
Microthelys rubrocallosa (Robinson & Greenman) Garay
BASIONYM: Spiranthes rubrocallosa Robinson & Greenman
SYNONYMS: Galeottiella rubrocallosa (Robinson & Greenman) Szlachetko
Schiedeella rubrocallosa (Robinson & Greenman) Burns-Balogh
Oeceoclades maculata (Lindley) Lindley*
BASIONYM: Angraecum maculatum Lindley
Pelexia adnata (Swartz) Sprengel
BASIONYM: Satyrium adnatum Swartz
SYNONYM: Spiranthes adnata (Swartz) Bentham in Fawcett
Phaius tankervilleae (Aiton) Blume
BASIONYM: Limodorum tankervilleae Aiton
Piperia candida R. Morgan & J. Ackerman
Platanthera candida (R. Morgan & J. Ackerman) R.M. Bateman nom. illeg.
Piperia colemanii Morgan & Glicenstein
SYNONYM: Platanthera colemanii (Morgan & Glicenstein) R.M. Bateman
Piperia cooperi (S. Watson) Rydberg
BASIONYM: Habenaria cooperi S. Watson
SYNONYM: Platanthera cooperi (S. Watson) R.M. Bateman
Piperia elegans (Lindley) Rydberg subsp. elegans
BASIONYM: Platanthera elegans Lindley
SYNONYMS: Habenaria 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
Platanthera unalascensis (Sprengel) Kurtz subsp. maritima (Greene) de Filipps

Piperia elongata Rydberg
SYNONYMS: 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
Piperia leptopetala Rydberg
SYNONYM: Platanthera leptopetala (Rydberg) R.M. Bateman
Piperia michaelii (Greene) Rydberg
BASIONYM: Habenaria michaelii Greene
SYNONYMS: Piperia elongata Rydberg subsp. michaelii (Greene) Ackerman
Platanthera michaelii (E. Greene) R.M. Bateman
Piperia transversa Suksdorf
SYNONYM: Platanthera transversa (Suksdorf) R.M. Bateman
Piperia unalascensis (Sprengel) Rydberg
BASIONYM: Spiranthes unalascensis Sprengel
SYNONYMS: Habenaria unalascensis (Sprengel) S. Watson
Platanthera unalascensis (Sprengel) Kurtz
MISAPPLIED NAME: Platanthera foetida Geyer in Hooker
Piperia yadonii R. Morgan & J. Ackerman
SYNONYMS: Platanthera yadonii (R. Morgan & J. Ackerman) R.M. Bateman
Habenaria californica Grinnell nom. nud.
Platanthera aquilonis Sheviak
MISAPPLIED NAMES: Habenaria 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
Platanthera blephariglottis (Willdenow) Lindley
BASIONYM: Orchis blephariglottis Willdenow
SYNONYMS: Blephariglottis blephariglottis (Willdenow) Rydberg
Habenaria blephariglottis (Willdenow) Hooker
Platanthera brevifolia (Greene) Kranzlein
BASIONYM: Habenaria brevifolia Greene
Platanthera chapmanii (Small) Luer emend. Folsom
BASIONYM: Blephariglottis chapmanii Small
SYNONYMS: Habenaria xchapmanii (Small) Ames
Platanthera xchapmanii (Small) Luer
Platanthera chorisiana (Chamisso) Reichenbach f.
BASIONYM: Habenaria chorisiana Chamisso
SYNONYMS: Limnorchis chorisiana (Chamisso) J.P. Anderson
Pseudodiphryllum chorisianum (Chamisso) Nevski
Platanthera ciliaris (Linnaeus) Lindley
BASIONYM: Orchis ciliaris Linnaeus
SYNONYMS: Blephariglottis ciliaris (Linnaeus) Rydberg
Habenaria ciliaris (Linnaeus) R. Brown
Platanthera conspicua (Nash) P.M. Brown
BASIONYM: Habenaria conspicua Nash
SYNONYMS: Blephariglottis conspicua (Nash) Small
Habenaria blephariglottis var. conspicua (Nash) Ames
Platanthera blephariglottis (Willdenow) Lindley var. conspicua (Nash) Luer
Platanthera convallariifolia Fischer in Lindley
SYNONYM: Limnorchis convallariaefolius (Fischer) Rydberg
114
MISAPPLIED NAME: Habenaria borealis Chamisso var. viridiflora Chamisso

Platanthera cristata (Michaux) Lindley
BASIONYM: Orchis cristata Michaux
SYNONYMS: Blephariglottis cristata (Michaux) Rafinesque
Habenaria cristata (Michaux) R. Brown
Platanthera dilatata (Pursh) Lindley
BASIONYM: Orchis dilatata Pursh
SYNONYMS: Limnorchis dilatata (Pursh) Rydberg in Britton
Piperia dilatata (Pursh) Szlachetko & P. Rutkowski
Platanthera dilatata (Pursh) Lindley var. angustifolia Hooker
Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour
BASIONYM: Habenaria borealis Chamisso var. albiflora Chamisso
SYNONYMS: Habenaria dilatata (Pursh) Hooker var. albiflora (Chamisso) Correll
Piperia dilatata (Pursh) Szlachetko & P. Rutkowski var. albiflora (Chamisso) Szlachetko & P. Rutkowski
Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer
BASIONYM: Platanthera leucostachys Lindley
SYNONYMS: Habenaria dilatata (Pursh) Hooker var. leucostachys (Lindley) Ames
Habenaria leucostachys (Lindley) S. Watson
Limnorchis leucostachys (Lindley) Rydberg
Platanthera flava (Linnaeus) Lindley var. flava
BASIONYM: Orchis flava Linnaeus
SYNONYMS: Habenaria flava (Linnaeus) R. Brown in Sprengel
Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer
BASIONYM: Habenaria herbiola R. Brown in Aiton
SYNONYMS: Habenaria flava (Linnaeus) R. Brown inSprengel 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
Platanthera grandiflora (Bigelow) Lindley
BASIONYM: Orchis grandiflora Bigelow
SYNONYMS: Blephariglottis grandiflora (Bigelow) Rydberg
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
Platanthera hookeri (Torrey) Lindley
BASIONYM: Habenaria hookeri Torrey
SYNONYMS: Habenaria hookeriana Torrey 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
Platanthera huronensis (Nuttall) Lindley
BASIONYM: Orchis huronensis Nuttall
SYNONYMS: Habenaria hyperborea (Linnaeus) R. Brown var. huronensis (Nuttall) Farwell
Habenaria xmedia (Rydberg) Niles
Limnorchis media Rydberg
115
Platanthera dilatata (Pursh) Lindley var. chlorantha Hultén

Platanthera hyperborea (Linnaeus) Lindley var. huronensis (Nuttall) Luer
Platanthera xmedia (Rydberg) Luer
Platanthera hyperborea (Linnaeus) Lindley
BASIONYM: Orchis hyperborea Linnaeus
SYNONYMS: Habenaria hyperborea (Linnaeus) R. Brown in Aiton
Limnorchis hyperborea (Linnaeus) Rydberg
Platanthera lacera (Michaux) G. Don
BASIONYM: Orchis lacera Michaux
SYNONYM: Fimbriella lacera (Michaux) F. Butzin
Habenaria lacera (Michaux) R. Brown
Platanthera leucophaea (Nuttall) Lindley
BASIONYM: Orchis leucophaea Nuttall
SYNONYMS: Fimbriella leucophaea (Nuttall) Farwell
Habenaria leucophaea (Nuttall) A. Gray
Platanthera limosa Lindley
SYNONYMS: Habenaria limosa (Lindley) Hemsley
Habenaria thurberi A. Gray
Limnorchis arizonica Rydberg
Platanthera macrophylla (Goldie) P.M. Brown
BASIONYM: Habenaria macrophylla Goldie
SYNONYMS: Habenaria orbiculata (Pursh) Hooker 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 obtusata (Banks in Pursh) Lindley
BASIONYM: Orchis obtusata Banks in Pursh
SYNONYMS: Habenaria obtusata (Banks in Pursh)Richards
Lysiella obtusata Rydberg
forma collectanea (Fernald) P.M. Brown
BASIONYM: Habenaria obtusata (Banks in Pursh) Richards var. collectanea Fernald
Platanthera orbiculata (Pursh) Lindley
BASIONYM: Orchis orbiculata Pursh
SYNONYMS: Habenaria orbiculata (Pursh) 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
Platanthera peramoena (A. Gray) A. Gray
BASIONYM: Habenaria peramoena A. Gray
SYNONYM: Blephariglottis peramoena (Gray) Rydberg
Fimbriella peramoena (A. Gray) F. Butzin
Platanthera praeclara Sheviak & Bowles
SYNONYMS: Fimbriella praeclara (Sheviak & M.L. Bowles) Szlachetko & Rutkowski
Habenaria leucophaea (Nuttall) A. Gray var. praeclara (Sheviak & Bowles) Cronquist
Platanthera psycodes (Linnaeus) Lindley
BASIONYM: Orchis psycodes Linnaeus
116
SYNONYMS: 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 purpurascens (Rydberg) Sheviak & W.F. Jennings
BASIONYM: Limnorchis purpurascens Rydberg
SYNONYM: Habenaria purpurascens (Rydberg) Tidestrom
Habenaria hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Ames
Platanthera hyperborea (Linnaeus) Lindley var. purpurascens (Rydberg) Luer
Platanthera sparsiflora (S. Watson) Schlechter
BASIONYM: Habenaria sparsiflora S. Watson
SYNONYMS: Limnorchis laxiflora Rydberg
Limnorchis sparsiflora (S. Watson) Rydberg
Platanthera stricta Lindley
SYNONYMS: 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
Platanthera tescamnis Sheviak & W.F. Jennings
SYNONYMS: Limnorchis ensifolia Rydberg
Platanthera sparsiflora (S. Watson) Schlechter var. ensifolia (Rydberg) Luer
Platanthera tipuloides (Linnaeus) Lindley var. behringiana (Rydberg) Hultén
BASIONYM: Limnorchis behringiana Rydberg
SYNONYM: Habenaria behringiana (Rydberg) Ames
Platanthera xandrewsii (Niles) Luer
BASIONYM: Habenaria xandrewsii M. White in G.G. Niles
SYNONYMS:: Blephariglottis xandrewsii (White) House
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
Platythelys latifolia (Linnaeus) Garay & Ormerod
BASIONYM: Satytium latifolium Linnaeus
SYNONYMS: Erythrodes sagreana (A. Richard) Leon
Physurus sagreanus A. Richard
117
Platythelys sagreana (A. Richard) Garay

Platythelys querceticola (Lindley) Garay
BASIONYM: Physurus querceticola Lindley
SYNONYM: Erythrodes querceticola (Lindley) Ames
Pleurothallis gelida Lindley
SYNONYM: Stelis gelida (Lindley) Pridgeon & M.W. Chase
Pogonia ophioglossoides (Linnaeus) Ker-Gawler
BASIONYM: Arethusa
ophioglossoides Linnaeus
forma brachypogon (Fernald) P.M. Brown
BASIONYM: Pogonia ophioglossoides (Linnaeus) Ker-Gawler var. brachypogon Fernald
Polystachya concreta (Jacquin) Garay & Sweet
BASIONYM: Epidendrum concretum Jacquin
MISAPPLIED NAME: Polystachya flavescens (Lindley) J. K. Small
Ponthieva brittoniae Ames
SYNONYM: Ponthieva racemosa (Walter) C. Mohr var. brittonae (Ames) Luer
Ponthieva racemosa (Walter) Mohr
BASIONYM: Arethusa racemosa Walter
Prescottia oligantha (Swartz) Lindley
BASIONYM: Cranichis oligantha Swartz
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 (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 pygmaea (Hooker) W.E. Higgins
BASIONYM: Epidendrum pygmaceum Hooker
SYNONYMS: Encyclia pygmaea (Hooker) Dressler
Hormidium pygmaeum (Hooker) Bentham & Hooker f.
Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E.
Higgins
BASIONYM: Epidendrum erythronioides Small
SYNONYMS: Encyclia bidentata (Lindley) Hágsater & 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
Pseudorchis straminea (Fernald) Soó
BASIONYM: Habenaria straminea Fernald
SYNONYMS: Habenaria albida (Linnaeus) 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
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
Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Sauleda, Wunderlin & Hansen
BASIONYM: Spiranthes lanceolata (Aublet) Leon var. paludicola Luer
118
SYNONYM: Stenorrhynchos cinnabarinms (La Llave & Lexarza) Lindley var. paludicola (Luer) W. J. Schrenk
Sacoila squamulosa (Kunth) Garay
BASIONYM: Neottia squamulosa Kunth
SYNONYMS: Sacoila lanceolata (Aublet)Garay var. squamulosa (Kunth) Szlachetko
Spiranthes squamulosa (Kunth) Leon
Stenorrhynchos squamulosum (Kunth) Sprengel
Schiedeella arizonica P.M. Brown
MISAPPLIED NAMES: Schiedeella fauci-sanguinea (Dod) Burns-Balogh
Schiedeella parasitica (A. Richard & Galeotti) Schlechter
Spiranthes fauci-sanguinea Dod
Spiranthes parasitica A. Richard & Galeotti
Spiranthes brevilabris Lindley
SYNONYM: Spiranthes gracilis (Bigelow) Beck var. brevilabris (Lindley) Correll
Spiranthes casei var. casei Catling & Cruise
MISAPPLIED NAME: Spiranthes intermedia Ames
Spiranthes cernua (Linnaeus) L.C. Richard
BASIONYM: Ophrys cernua Linnaeus
SYNONYMS: Ibidium cernuum (Linnaeus) House
Triorchis cernua (Linnaeus) Nieuwland
Spiranthes delitescens Sheviak
MISAPPLIED NAME: Spiranthes graminea Lindley
Spiranthes diluvialis Sheviak
SYNONYM: Spiranthes romanzoffiana Chamisso var. diluvialis (Sheviak) S.L. Welsh
Spiranthes floridana (Wherry) Cory emend. P.M. Brown
BASIONYM: Ibidium floridanum Wherry
SYNONYMS: Spiranthes brevilabris Lindley var. floridana (Wherry) Luer
Spiranthes gracilis var. floridana (Wherry) Correll
Spiranthes lacera Rafinesque var. lacera
BASIONYM: Neottia lacera Rafinesque
Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer
BASIONYM: Neottia gracilis Bigelow
SYNONYMS: Ibidium beckii (Lindley) House
Ibidium gracile (Bigelow) House
Spiranthes beckii Lindley
Spiranthes gracilis (Bigelow) Beck
Spiranthes laciniata (Small) Ames
BASIONYM: Gyrostachys laciniata Small
SYNONYM: Ibidium laciniatum (Small) House
Triorchis laciniata House
Spiranthes longilabris Lindley
SYNONYMS: Ibidium longilabre (Lindley) House
Spiranthes brevifolia Chapman
Triorchis longilabris House
Spiranthes lucida (H.H. Eaton) Ames
BASIONYM: Neottia lucida H.H. Eaton
SYNONYMS: Ibidium plantagineum (Rafinesque) House
Spiranthes plantaginea Rafinesque
Triorchis plantaginea Nieuwland
Spiranthes ochroleuca (Rydberg) Rydberg
BASIONYM: Gyrostachys ochroleuca Rydberg in Britton
SYNONYMS: Spiranthes cernua (Linnaeus) L.C. Richard var. ochroleuca (Rydberg) Ames
Spiranthes xsteigeri Correll
Triorchis cernua Nieuwland var. ochroleuca Farwell
Triorchis ochroleuca Nieuwland
119
Spiranthes odorata (Nuttall) Lindley

BASIONYM: Neottia odorata Nuttall
SYNONYMS: Ibidium odoratum (Nuttall) House
Spiranthes cernua (Linnaeus) L.C. Richard var. odorata (Nuttall) Correll
Triorchis odorata (Nuttall) Nieuwland
Spiranthes ovalis Lindley
SYNONYM: Ibidium ovale (Lindley) House
Spiranthes porrifolia Lindley
SYNONYMS: Gyrostachys porrifolia (Lindley) Kuntze
Ibidium porrifolium Rydberg
Spiranthes romanzoffiana Chamisso var. porrifolia (Lindley) Ames & Correll
Spiranthes praecox (Walter) S. Watson
BASIONYM: Limodorum praecox Walter
SYNONYM: Ibidium praecox (Walter) House
Triorchis praecox (Walter) Nieuwland
Spiranthes romanzoffiana Chamisso
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 torta (Thunberg) Garay & Sweet
BASIONYM: Ophrys torta Thunberg
SYNONYM: Ibidium tortile (Swartz) House
MISAPPLIED NAME: Spiranthes tortilis (Swartz) Richard
Spiranthes tuberosa Rafinesque
SYNONYMS: Ibidium beckii House
Spiranthes grayi Ames
Spiranthes tuberosa var. grayi (Ames) Fernald
Triorchis grayi (Ames) Nieuwland
Spiranthes vernalis Engelmann & A. Gray
SYNONYMS: 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, Sterns & Poggenberg
Tolumnia bahamense (Nash in Britton & Millspaugh) G. J. Braem
BASIONYM: Oncidium bahamense Nash in Britton & Millspaugh
MISAPPLIED NAME: Tolumnia variegata (Swartz) G. J. Braem
Trichocentrum carthagenense (Jacquin) M.W. Chase & N.H. Williams
BASIONYM: Epidendrum carthagenense Jacquin
SYNONYM: Lophiaris carthagenensis (Jacquin) Braem
Trichocentrum luridum (Lindley) M.W. Chase & N.H. Williams
BASIONYM: Oncidium luridum Lindley
SYNONYM: Lophiaris lurida (Lindley) Braem
Trichocentrum undulatum (Swartz) Ackerman & Chase
BASIONYM: Epidendrum undulatum Swartz
SYNONYMS: Oncidium undulatum (Swartz) Salisbury
Trichocentrum maculatum (Aublet) M. Chase & N.A. Williams nom. illeg.
120
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
Triphora amazonica Schlechter
SYNONYM: Triphora latifolia G. Luer
Triphora gentianoides (Swartz) Ames & Schlechter*
BASIONYM: Limodorum gentianoides Swartz
SYNONYMS: Pogonia cubensis Reichenbach f.
Pogonia gentianoides (Swartz) Sprengel
Triphora cubensis (Reichenbach f.) Ames
Triphora rickettii Luer
MISAPPLIED NAME: Triphora yucatanensis Ames
Triphora trianthophoros (Swartz) Rydberg
BASIONYM: Arethusa trianthophoros Swartz
SYNONYMS: Arethusa pendula Mühlenberg in Willdenow
Pogonia pendula (Mühlenberg in Willdenow) Lindley
Pogonia trianthophoros (Swartz) Britton, Sterns & Poggenberg
Triphora pendula (Mühlenberg in Willdenow) Nuttall
Tropidia polystachya (Swartz) Ames
BASIONYM: Serapias polystachya Swartz
SYNONYM: Tropidia eatonii Ames
Vanilla barbellata Reichenbach f.
SYNONYM: Vanilla articulata Northrop
Vanilla dilloniana Correll
MISAPPLIED NAME: Vanilla eggersii Rolfe
Vanilla inodora Schiede
SYNONYMS: Epidendrum vanilla Linnaeus p.p.
Vanilla mexicana Miller
Vanilla planifolia Andrews
SYNONYM: 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
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. islandicum (Lindley) Schulze = Coeloglossum viride
(Linnaeus) Hartman var. viride
122
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
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.
Dactylorhiza praetermissa (Druce) Soó var. junialis (Vermeulen) Senghas = Dactylorhiza majalis subsp.
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
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
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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
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
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
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 =
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
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
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
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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
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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
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
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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
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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
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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
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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
Clamshell
orchid,
Prosthechea
cochleata var.
triandra
140
crested
coralroot
Hexalectris
spicata
141
ghost orchid
Dendrophylax
lindenii
all
photographs
© Connie
Bransilver
Barrow
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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,
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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
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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
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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
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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
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 myco-
heterotrophic 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
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
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McGrath: PLATANTHERA PALLIDA

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 YELLOW-

CRESTED 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).
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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).
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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 Correll Range Sag Harbor Napeague Montauk
Lip width ---- 2 2 2

Dorsal sepal length 3-5 4 4 4
Dorsal sepal width 2-3 2.5 4 2.4
Lateral petal length 2-4 3 3* 3
Lateral petal width 2-4 3 2.5* 3
Lateral sepal length 3-4 3 3 4
Lateral sepal width 2-3 3 2.75 2.5
Spur length 5-10 6 6 ----
* when present
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
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
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
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
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 over-
population 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
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
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
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
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 white-
flowering 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
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
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
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 near-
xeric 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
3 4
Platanthera pallida
5 5 6
Platanthera cristata forma straminea
165
center: Platanthera xcanbyi

right: Platanthera blephariglottis
87 7 8
5
Platanthera cristata
center: Platanthera xbeckneri

right: Platanthera conspicua 9 10
166
center: Platanthera xbicolor

right:: Platanthera blephariglottis
12 8
11
Platanthera ciliaris
center: Platanthera xlueri

13 10
right: Platanthera conspicua
167
Ferry: STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS
STOMATA, SUBSIDIARY CELLS, AND IMPLICATIONS
Robert J. Ferry
Reprinted with permission from McAllen International Orchid Society Journal 9(3) 9-16. 2008
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
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
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, Fig. 2. Abaxial epidermal leaf cells, x80,
Stanhopea oculata. Pl#290496-5 C6. Xylol strip Abα- Stanhopea oculata. Pl#290496-5 C6. Xylol strip Abα-2.
2. 26 May, 1996. 35mm Photomicrograph 26 May, 1996. 35mm Photomicrograph transparency
transparency 19 July, 1996.
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
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. Fig. 5. Paracytic Subsidiary cells. Drawing,

Drawing, 01 February, 2008. 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
Fig. 6. Diacytic Subsidiary cells. Drawing,
01 February, 2008. 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
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.
RECENT WORK
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.
Fig. 8. Abaxial epidermal leaf cells, S. stellata. Dried FIG. 9 Abaxial epidermal leaf cells, S. stellata. Dried
leaf, X80 mag, 35mm transp#5, 19Jan08. leaf, X320 mag, 35mm transp#7, 19Jan08
172
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. FIG. 11 Abaxial epidermal leaf cells, S. cernua. Live leaf,
Dried leaf, X80 mag, 35mm transp#16, 19Jan08 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
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 non-
flowering 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
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
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 wide-
ranging 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,
Intermediate in characters between the two parents Calopogon barbatus and Calopogon multiflorus
177
Type: U.S.A. Florida: Wakulla Co., Ochlockonee River State Park. 10 April 2005 P.M. Brown 05-
14 (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,
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,
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,
Intermediate in characters between the two parents Calopogon barbatus and Calopogon pallidus
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
Calopogon xgoethensis P.M. Brown nothosp. nov

Planta inter Calopogon multiflorus et Calopogon tuberosus intermedia et habitu, colore et forma florum,
Intermediate in characters between the two parents Calopogon multiflorus and Calopogon tuberosus
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 C. multiflorus
Figure 1. Calopogon hybrids and parentage
179
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
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 C.. tuberosus

181
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
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183
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184

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

NOTES FROM THE EDITOR

Paul Martin Brown

ORCHID FLORA OF THE FLORIDA PANTHER NATIONAL WILDLIFE

In an effort to conserve the native orchid diversity present on the FPNWR, a

NATURAL COMMUNITIES OF THE FPNWR

A typical hardwood hammock on the FPNWR. Slash pine (Pinus elliotti) is

A typical pine flatwood on the FPNWR.

Sloughs and Strand Swamps

Scientific name: Bletia purpurea (Lamark) de Candolle

Scientific name: Calopogon multiflorus Lindley

Typical rose-pink color. Forma albiflorus from the FPNWR.

Scientific name: Campylocentrum pachyrrhizum (Reichenbach f.) Rolfe

Scientific name: Cyrtopodium punctatum (Linnaeus) Lindley

Scientific name: Dendrophylax lindenii (Lindley) Bentham ex Rolfe

Note one open flower and two buds.

Scientific name: Encyclia tampensis (Lindley) Small

Scientific name: Epidendrum amphistomum A. Richard

Scientific name: Epidendrum floridense Hágsater

Scientific name: Epidendrum nocturnum Jacquin

Scientific name: Epidendrum rigidum Jacquin

Scientific name: Eulophia alta (Linnaeus) Fawcett & Rendle

Vegetative plants with typical palm-like leaves.

Scientific name: Habenaria odontopetala Reichenbach f.

Scientific name: Habenaria quinqueseta (Michaux) Eaton

Scientific name: Habenaria repens Nuttall

Scientific name: Harrisella porrecta (Reichenbach f.) Fawcett & Rendle

Scientific name: Ionopsis utricularioides (Swartz) Lindley

Scientific name: Oeceoclades maculata (Lindley) Lindley

Scientific name: Polystachya concreta (Jacquin) Garay & Sweet

Scientific name: Sacoila lanceolata (Aublet) Garay var. lanceolata

Typical red-coral color. Forma albidaviridis in capsule.

Scientific name: Spiranthes longilabris Lindley

Scientific name: Spiranthes odorata (Nuttall) Lindley

Typical yellow-throated color type. Green-throated color type.

Scientific name: Spiranthes vernalis Engelmann & Gray

Scientific name: Zeuxine strateumatica (Linnaeus) Schlechter

Species from Adjacent Lands

1) Bulbophyllum pachyrhachis – hardwood hammocks, sloughs, and strand swamps

Scott L. Stewart, Ph.D. Larry W. Richardson

SYNONYMS AND MISAPPLIED NAMES OF WILD ORCHIDS

Amerorchis rotundifolia (Banks) Hultén

forma pallidum (House) Barnhart

Cleistes bifaria (Fernald) Catling & Gregg

Corallorhiza unguiculata Rafinesque

SYNONYM: Epidendrum conopseum R. Brown

Govenia floridana P.M. Brown

BASIONYM: Arethusa medeoloides Pursh

SYNONYM: Hammarbya paludosa Kuntze

Platanthera unalascensis (Sprengel) Kurtz subsp. maritima (Greene) de Filipps

MISAPPLIED NAME: Habenaria borealis Chamisso var. viridiflora Chamisso

Platanthera dilatata (Pursh) Lindley var. chlorantha Hultén

SYNONYMS: Blephariglottis psycodes (Linnaeus) Rydberg

Platythelys sagreana (A. Richard) Garay

Spiranthes odorata (Nuttall) Lindley

forma flavovirens (P.M. Brown) P.M. Brown

CROSS-REFERENCES FOR SYNONYMS AND MISAPPLIED NAMES

Arethusa divaricata Linnaeus = Cleistes divaricata (Linnaeus) Ames

Cypripedium occidentale Watson = Cypripedium montanum Douglas in Lindley

Habenaria brevifolia Greene = Platanthera brevifolia (Greene) Kranzlein

Habenaria leucophaea (Nuttall) A. Gray = Platanthera leucophaea (Nuttall) Lindley

Vanilla fragrans (Salisbury) Ames = Vanilla planifolia Andrews