June 2009 North American Native Orchid Journal

NORTH AMERICAN
NATIVE ORCHID JOURNAL

Volume 15(2) 2009
IN THIS ISSUE:
RECENT PUBLICATIONS REGARDING PLATANTHERA AND ALLIED GENERA
THE AMERICAN SPECIES OF LIMNORCHIS
…OF CABBAGES AND KINGS
PLATANTHERA ×CANBYI ON THE EASTERN SHORE OF MARYLAND
and more………….
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.
NORTH AMERICAN
NATIVE ORCHID JOURNAL
Volume 15 (2) 2009
CONTENTS
NOTES FROM THE EDITORS
64
AN IMPORTANT NOTE
64
RECENT PUBLICATIONS REGARDING PLATANTHERA AND ALLIED GENERA,
NEW COMBINATIONS, AND SOME NOVEL OBSERVATIONS
Paul Martin Brown, Scott L. Stewart and authors
65
Per Axel Rydberg
a facsimile reprint
88
WILD ORCHIDS IN THE 21ST CENTURY:
field guides and floras on North American native orchids
2007-2009 update
117
…OF CABBAGES AND KINGS
The Slow Empiricist
122
RECENT ARTICLES OF INTEREST IN OTHER JOURNALS
125
REDISCOVERIES OF PLATANTHERA ×CANBYI ON THE
EASTERN SHORE OF MARYLAND
Chelsea Kieffer
128
BOOK REVIEWS
130
NEW COMBINATIONS IN VOLUME 15(2)
132
Unless otherwise credited, all drawings in this issue are by Stan Folsom and photographs by P.M. Brown.
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 15(2): 64-137 issued September 22, 2009.
Copyright 2009 by the North American Native Orchid Journal
Cover: Platanthera (Blephariglottis) ciliaris by Stan Folsom
NOTES FROM THE EDITORS
The second issue of the North American Native Orchid Journal for 2009 presents a
digest version of three recent papers and their proposals as well as some new
combination to go with them. It has been an interesting process working with three
publishers, sets of authors, and completing the translation form the German. Two
more issues for 2009 are being planned. In late November A Tale of Two Trips will
present two lengthy articles about orchid hunting in the western states and an exciting
summer of orchid discoveries for the Senior Editor will result in the December issue
being devoted entirely to the wild orchids of the White Mountains of New Hampshire
and nearby Maine.
The electronic format continues to be well received and we now reach more
than 1500 readers.
You may read back issues at: http://wiki.terrorchid.org/tow:journals. The
current update of the North American Personal Checklist is also available at that website.
The checklist will be updated as needed with new taxa and changes noted.
Paul Martin Brown, Editor

naorchid@aol.com
10896 SW 90th Terrace, Ocala, FL 34481
36 Avenue F, Acton, Maine 04001 (June- early October)
Scott L. Stewart, PhD. Associate Editor

slstewar@gmail.com
Kankakee Community College
Horticulture & Agriculture Programs
100 College Drive
Kankakee, Illinois 60901
AN IMPORTANT NOTE
In the description of Cleistesiopsis oracamporum NANOJ 15(1): 52-53
several changes have been made since the original posting. All print
copies were corrected and the current on-line posting is correct.
Please note the change in spelling from oracamporum to
oricamporum and the Latin description has been greatly improved
thanks to work by Dr. K. Gandhi, Harvard University Herbaria.
Other minor corrections or changes were made a few other places in
that issue as well.
PMB
Brown, Stewart, et al.: RECENT PUBLICATIONS REGARDING PLATANTHERA
RECENT PUBLICATIONS REGARDING PLATANTHERA AND ALLIED

GENERA, NEW COMBINATIONS, AND SOME NOVEL OBSERVATIONS
or
Oh, No, Here They Go Again
Paul Martin Brown, Scott L. Stewart and authors
In the past months three papers of interest to North American native orchid enthusiasts have appeared in three
different publications. Each dealt with generic limits within Platanthera and its allies and presented different points of
view and often resulting new combinations. One of the goals of the North American Native Orchid Journal is to bring
current taxonomic views to our readers. The following are selected quotes and are NOT complete texts and are included
here with the permission of the respective editors of those journals; those who wish to learn more concerning these
topics or view the extensive graphics are urged to access the original publications. Perhaps the most important thing to
remember is that much of this falls under ‗taxonomic opinion‘. Apart from correcting errors, especial regarding priority
of names - an example in another genus would be Listera banksiana Lindley 1840 versus the more widely used Listera
caurina Piper 1898 - genus, and often species, concepts can be a very personal choice. Which genus you chose to
recognize is often a matter of preference, apart from those that are more or less universally accepted. A good example is
that Habenaria and Platanthera s.l. are very different from each other and not even that closely related. Whether or not you
regard Platanthera as a large umbrella genus or prefer Piperia, Gymnadeniopsis, Limnorchis, etc. is a choice you may make.
Some generic changes are very clear-cut and well-accepted in the molecular world but slow to take in the realm of the
native orchid enthusiast, even though many of these ‗changes‘ hie back to earlier, and often original, names. Regarding
species, because of the International Rules of Nomenclature, certain species epithets must be used with certain genera,
although most are transferable. And, of course, each author of new species/varieties and new combination feels strongly
that they are correct! Genera and species chosen for the widely accessible checklists are again the opinion of individuals
or working groups, such as the Monocot Checklist, World Checklist, USDA, NANOJ checklist, etc. There is really no
such thing as a universally accepted name apart from the correct name regarding publication and priority. Table and
figure numbers refer to the original publications and bibliographical references are included for only the extractions. A
glossary of terms, particularly relating to molecular work, is included at the end of the compiloarticle.
MOLECULAR PHYLOGENETICS AND MORPHOLOGICAL REAPPRAISAL OF THE

PLATANTHERA CLADE (ORCHIDACEAE: ORCHIDINAE) PROMPTS EXPANSION OF
THE GENERIC LIMITS OF GALEARIS AND PLATANTHERA
Richard M. Bateman, Karen E. James, Yi-Bo Luo, Robert K. Lauri, Timothy Fulcher,
Phillip J. Cribb, and Mark W. Chase
Originally published in Annals of Botany 104: 431-445. 2009
Reprinted by permission of publisher.
 Background and Aims: The Platanthera clade dominates the North American orchid flora and is well
represented in eastern Asia. It has also generated some classic studies of speciation in Platanthera sections
Platanthera and Limnorchis. However, it has proved rich in taxonomic controversy and near-monotypic genera.
The clade is reviewed via a new molecular phylogenetic analysis and those results are combined with brief
65
reconsideration of morphology in the group, aiming to rationalize the species into a smaller number of larger
monophyletic genera and sections.
 Methods: Nuclear ribosomal internal transcribed spacer (ITS) sequences were obtained from 86 accessions of
35 named taxa, supplemented from GenBank with five accessions encompassing a further two named taxa.
 Key Results: Using Pseudorchis as outgroup, and scoring indels, the data matrix generated 30 most-parsimonious
trees that differed in the placement of two major groups plus two closely related species. Several other internal
nodes also attracted only indifferent statistical support. Nonetheless, by combining implicit assessment of
morphological divergence with explicit assessment of molecular divergence (when available), nine former
genera can be rationalized into four revised genera by sinking the monotypic Amerorchis, together with
Aceratorchis and Chondradenia (neither yet sequenced), into Galearis, and by amalgamating Piperia, Diphylax and the
monotypic Tsaiorchis into the former Platanthera section Platanthera. After further species sampling, this section
will require sub-division into at least three sections. The present nomenclatural adjustments prompt five new
combinations.
 Conclusions: Resolution of major groups should facilitate future species-level research on the Platanthera clade.
Recent evidence suggests that ITS sequence divergence characterizes most species other than the P. bifolia
group. The floral differences that distinguished Piperia, Diphylax and Tsaiorchis from Platanthera, and Aceratorchis
and Chondradenia from Galearis, reflect various forms of heterochrony (notably paedomorphosis); this affected
both the perianth and the gynostemium, and may have proved adaptive in montane habitats. Floral reduction
was combined with lateral expansion of the root tubers in Piperia and Diphylax (including Tsaiorchis), whereas
root tubers were minimized in the putative (but currently poorly supported) Neolindleya–Galearis clade.
Allopolyploidy and/or autogamy strongly influenced speciation in Platanthera section Limnorchis and perhaps
also Neolindleya. Reproductive biology remains an important driver of evolution in the clade, though plant–
pollinator specificity and distinctness of the species boundaries have often been exaggerated.
INTRODUCTION
Platanthera Rich. (the butterfly orchids) is one of the most species-rich genera of North
Temperate orchids (Hapeman and Inoue, 1997), having centres of diversity in East Asia (Inoue,
1983) and North America (Luer, 1975; Sheviak, 2002). Platanthera and allied genera share with many
other temperate orchid groups a long and complex taxonomic history. Victorian luminaries
including Robert Brown and Joseph Hooker lumped Platanthera species into Habenaria, misled by
superficial floral similarities of these genera evident in, for example, rostellum morphology (cf.
Dressler, 1993). Other authors elected to divide Platanthera into several genera, each of which
typically contained few (1–5) species. During the second half of the 20th century, the view most
commonly expressed by morphological systematists recognized the majority of these former genera
as sections of the genus Platanthera (e.g. Lysiella, Limnorchis, Blephariglottis and Tulotis). In contrast,
other relevant taxa became more commonly recognized as distinct genera, notably Pseudorchis Seguier
(syn. Leucorchis A. & D. Löve), Piperia Rydb. (1901a) and Diphylax Hook. f. (1869), together with the
more obscure, monotypic Tsaiorchis Tang & Wang (1936).
The most notorious ‗taxonomic football‘ was Piperia, which was over the last decade, these
intuitively inferred boundaries of, and relationships among, genera have been systematically revised
(and, in many cases, overturned) by sequence-based phylogenetic analyses using the rapidly evolving
nuclear ribosomal internal transcribed spacers (ITS) of nuclear ribosomal DNA (nrDNA). Pridgeon
et al. (1997) and Bateman et al. (1997) identified a weakly supported Platanthera clade, consisting of
Pseudorchis (one species analysed) as sister to Galearis (two species) plus Platanthera (six species, acting
as placeholders for putative sections within this larger genus, estimated at .85 species by Hapeman
and Inoue, 1997). These generic relationships were also obtained (again with limited statistical
support) by Bateman et al. (2003), in a study benefiting from substantially increased taxon sampling.
Pseudorchis (two species) was again shown as basally divergent, followed by the monotypic Amerorchis,
which was placed as sister to Galearis (three species). The next branch showed the monotypic
66
Neolindleya as sister to Platanthera sensu stricto (s.s.). Lastly, the genus Platanthera proved to encompass
several former genera that were already incorporated within Platanthera by most authorities
(represented by eight species). However, also nested within the genus was a strongly supported
monophyletic group of the three sampled species of Piperia. Consequently, all ten Piperia species
recognized by Ackerman and Morgan (2002) were duly incorporated in Platanthera as a new section
of the genus (Table 4 of Bateman et al., 2003).
Delimitation and relationships of genera and sections in the Platanthera clade

The Platanthera sensu lato (s.l.) clade is undoubtedly sister to Dactylorhiza s.l. plus Gymnadenia s.l.,
and is far more distantly related to Habenaria, despite the similar rostellar morphologies of the two
genera (Bateman et al., 2003). With the notable exception of Pseudorchis, mutation rate appears to be
substantially higher in the Platanthera clade than in its sister group, the Dactylorhiza s.l.–Gymnadenia s.l.
clade (fig. 1c of Box et al., 2008). Within the Platanthera clade, it remains unclear whether Pseudorchis
or the Galearis clade is sister to Platanthera s.s., but it is clear that Pseudorchis merits the status of genus
(contra Luer, 1975; Delforge, 2006). The two ITS branches that define the dichotomy between the
Galearis clade (tentatively including Neolindleya) and Platanthera s.s. are only weakly supported, despite
the fact that the two clades are readily separated using morphology.
Characters listed by Hapeman and Inoue (1997) as delimiting the genus Platanthera included
its broad anther, undivided stigma and fusiform tubers, though reports of exceptions for each
character have increased in number as the former genera Piperia and Diphylax have successively been
incorporated into the genus (Bateman et al., 2003; present study; also Lauri, 2007, who disagrees
with this degree of ‗lumping‘ at the genus level).
Within Platanthera, most of the sections recognized by Hapeman and Inoue (1997) survive
the present analysis. However, the monophyly of section Platanthera sensu Hapeman and Inoue (1997)
is challenged by the fact that the former genus Diphylax (and possibly also the former genus Piperia)
is embedded deeply within it, and by the placement of P. orbiculata (attributed to section Platanthera
by Hapeman and Inoue, 1997) as sister to P. oligantha of section Lysiella (Fig. 1). Admittedly, this
placement is decidedly tentative. Several experiments with taxon and character sampling separated P.
orbiculata from P. oligantha and placed it further down the tree. Also, P. orbiculata and P. obtusata
(putatively a close relative of P. oligantha) were separated (albeit both occurring within section
Platanthera) in the analyses of Hapeman and Inoue (1997). Lastly, P. orbiculata and P. oligantha are
subtended by the longest branches in our tree, thus raising the possibility that their apparent sister-
group status may reflect long-branch attraction.
Irrespective of their mutual relationships, valid cases can be made for recognizing the former
genera Lysiella (here represented by P. oligantha) and Lysias (here represented by P. orbiculata) as
sections of Platanthera (Fig. 1). The assignment of P. orbiculata to section Platanthera by Hapeman and
Inoue (1997) is not supported.
The Galearis clade: Galearis s.l.

The taxonomic challenges presented within the Galearis clade are analogous to those
previously presented by the Eurasian Himantoglossum clade (Bateman et al., 2003), in which a core
67
group of several species of Himantoglossum s.s. had as its unambiguous sister the possibly monotypic,
morphologically similar Barlia. This clade in turn had as its unambiguous sister the unequivocally
monotypic, superficially morphologically distinct Comperia, and this clade in turn had as its
ambiguous sister the unequivocally monotypic, more substantially morphologically distinct Steveniella.
Each former genus was separated by a moderate degree of disparity in ITS sequences. Thus,
Bateman et al.(2003) chose to sink both Barlia and Comperia into Himantoglossum, but retained
Steveniella as a separate genus Bateman et al. because of its less confident phylogenetic placement and
its comparatively distinct morphology (small, purple-washed leaves, central lobe of the labellum not
further divided, spur apex clearly bifid, gynostemium compact).
In the case of the Galearis clade, Amerorchis mimics the role of Comperia and Neolindleya
mimics the role of Steveniella. Both genera are commonly viewed as monotypic. Amerorchis is
undoubted sister to Galearis s.s., though they are separated by a substantial molecular distance.
However, comparison of the morphological descriptions of the two genera given in recent
publications (e.g. Wood and Neiland, 2001a, b; Sheviak and Catling, 2002) showed the two supposed
genera to be similar in most characters. Moreover, characters that do show some difference are
those that are known to be highly homoplastic within subtribe Orchidinae (Bateman et al., 2003),
notably Amerorchis more often having only one expanded leaf rather than two, and tending to have a
more clearly three-lobed labellum. Gynostemium morphology differs between Amerorchis, North
American Galearis s.s. (i.e. G. spectabilis, the type species of the genus) and southeast Asian Galearis s.s.
(Luo and Chen, 2000; Perner and Luo, 2007; R. Lauri, unpubl. res.). Amerorchis possesses an
undivided bilobed bursicle and G. spectabilis a divided bursicle, whereas Asian Galearis lack a bursicle
(Perner and Luo, 2007). The stigma is divided into two zones separated by the rostellum in both
Amerorchis (R. Lauri, unpubl. res.) and southeast Asian Galearis s.s. (Perner and Luo, 2007), whereas it
delineates a single inverted triangle in North American Galearis s.s. In addition, Amerorchis rotundifolia
has a distribution in North America that largely complements that of the more southerly Galearis
spectabilis, exhibiting only a narrow zone of overlap.
Their successive divergence below the two Asian species of Galearis in Fig. 1 implies that the
Asian species are derived relative to the North American species (Bateman et al., 2003). Considered
together, these factors arguably suggest that Amerorchis is better incorporated into Galearis than
retained as a monotypic genus. The relevant combination is made here in Table 1; surprisingly, this
species has not previously been placed in Rafinesque‘s (1833) Galearis, but rather was variously
assigned to Habenaria, Orchis and (more credibly) Platanthera, before being isolated as Amerorchis as
recently as 1968 (Hultén, 1968). Luer (1975) and Sheviak and Catling (2002) recognized only one
North American and one east Asian species of Galearis (excluding Amerorchis), whereas Wood and
Neiland (2001b) suggested 3–6 species in total, Cribb and Gale (2009) listed five species (including
the former genus Aceratorchis: see below) and Perner and Luo (2007) recognized six species in China
alone. Expanding his geographical coverage, Hunt (1971) listed 11 species in Asia alone (including
Aorchis sensu Vermeulen, 1972). Our analysis shows substantial molecular divergence between the
two East Asian species analysed, G. cyclochila and G. diantha, both of which appear morphologically
distinct from G. wardii (illustrated from China as ‗Orchis’ wardii by Chen et al., 1999, p. 306). We
suspect that there are substantially more than two bona fide species in the genus Galearis.
In addition, after a decade of failing to obtain DNA samples of the rare Chinese species
Aceratorchis tschilensis Schlechter (1922), we have nonetheless taken the radical step of incorporating
Aceratorchis into Galearis, an action suggested tentatively by Cribb and Wood (2001) and more firmly
68
by Cribb and Gale (2009). [Perner and Luo (2007, p. 135)] went one step further in suggesting that
‗A.’ tschiliensis is conspecific with Galearis roborowskii.
Our decision to place Aceratorchis within Galearis is based partly on similarities in vegetative
and gynostemium characters and partly on the hypothesis that the near radial symmetry of the
perianth of Aceratorchis and its spurless condition reflect an origin from ‗wild-type‘ Galearis flowers
via a heterochronic developmental shift termed pseudopeloria (Rudall and Bateman, 2002; Bateman
et al., 2003; Bateman and Rudall, 2006; Mondragón-Palomino and Theissen, 2008, 2009). If so, they
constitute atavistic reversals, rather than showing the fundamentally primitive morphology inferred
by Chen (1982) and Chen and Tsi (1987).
The Platanthera clade: section Limnorchis

In the analysis of Bateman et al. (2003), section Limnorchis was represented by only a single
placeholder, P. (hyperborea) viridiflora. Here, the 11 sequences of Platanthera section Limnorchis
generated by us or by Wallace (2002) (Appendix) show moderate divergence and constitute a well
supported monophyletic group. Indeed, the morphological distinction between section Limnorchis
and section Platanthera was recently reinforced by Gamarra et al. (2008), who documented
significantly different seed types in the two sections. Our 11 accessions supposedly represent five
species, and the five accessions of P. dilatata (all subtly molecularly different) represent three varieties
sensu Luer (1975) and Sheviak (2002). The samples are resolved into three molecularly distinct
groups, based respectively on P. stricta, P. dilatata and P. sparsiflora (Fig. 1). Despite their similarity in
morphology (noted by Luer, 1975), we originally hypothesized that the substantial divergence
between the dilatata (originally referred to hyperborea) and sparsiflora groups reflected contrasting
geographical foci; the former characterizes northwestern North America, whereas the latter, which
appeared relatively derived, is concentrated in the uplands of Mexico and southwestern North
America. However, the subsequent inclusion in our analysis of two accessions of P. aquilonis, a
species widespread in temperate North America (Sheviak, 1999), may challenge this interpretation.
The accession assigned to P. aquilonis by L. Wallace was placed firmly in the southerly sparsiflora
group, whereas the Alaskan accession tentatively assigned to P. aquilonis by R. Lauri was placed
within the northerly P. dilatata group (Fig. 1). Further molecular and morphological study of this
species group is therefore required.
The three accessions of the sparsiflora group analysed show moderate levels of both sequence
divergence (Fig. 1) and morphological divergence (Sheviak, 2002). If correctly identified, the Sedona
population of P. limosa found by us represents a small northward extension of the dominantly
Mexican range given by Luer (1975). Morphologically, it is differentiated from the similarly but more
widely distributed P. sparsiflora by its smaller flowers, narrower gynostemium and a labellum bearing
a comparatively long spur and a small basal callus convergent with that commonly found in section
Tulotis (Luer, 1975; Hapeman and Inoue, 1997; Sheviak, 2002). Platanthera aquilonis is facultatively
autogamous (Wallace, 2006) and has a dorsiventrally compressed gynostemium; previous
morphological comparisons with P. hyperborea s.s. are contradicted by the placement of Wallace‘s
sample but encouraged by the placement of Lauri‘s sample. Wallace (2003) observed approximately
equal separation of P. sparsiflora, P. aquilonis, P. dilatata and P. stricta in a population-level analysis that
combined inter simple sequence repeats (ISSRs) and randomly amplified polymorphic DNAs
(RAPDs).
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Because of the exceptional taxonomic instability surrounding the delimitation of P. hyperborea

and its putative allies (here represented by P. viridiflora and P. dilatata s.l.: cf. Luer, 1975; Wood and
Neiland, 2001c; Sheviak, 2002), we have chosen to refer to the sister of the sparsiflora group as the
dilatata group (Figs. 1 and 2). Bona fide species of the dilatata group have distributions that stretch
from the Aleutians in the northwest southwards to California and eastwards through Canada and
northeastern USA to Greenland, whereas P. hyperborea stretches to Iceland (Luer, 1975; Sheviak,
2002). Luer (1975) awarded viridiflora only varietal status under P. hyperborea, arguing that viridiflora
was merely a more robust form of P. hyperborea with a longer labellum and a longer spur (greater
than, as opposed to less than, 6 mm). However, Hapeman and Inoue (1997) revealed considerable
ITS divergence between P. hyperborea and P. viridiflora, which did not prove to be sisters, thus strongly
suggesting that viridiflora merits species-level distinction. It includes the western-most populations
commonly attributed to the P. hyperborea group, occurring in Alaska, the Aleutians and Kamchatka
(Luer, 1975; Sheviak, 2002). Supposed outlying populations of P. viridiflora in Hawaii have since been
referred to P. holochila; they appear to be relatively distantly related to P. viridiflora within section
Limnorchis (R. Lauri, unpubl. res.).
Certainly, the substantial divergence of P. stricta from the P. dilatata group challenges
Sheviak‘s (2002) synonymization of P. viridiflora into P. stricta. Considerable molecular divergence
between these species was also reported by Hapeman and Inoue (1997), though admittedly in our
analysis it costs only one additional step to place P. stricta as sister to the P. dilatata group, despite the
seven-step length of the branch separating P. stricta from the P. dilatata group (Fig. 1). The
distribution of P. stricta coincides with those of P. dilatata and P. hyperborea, stretching from the
Aleutians to northern California and Nevada. Morphologically similar to P. dilatata and P. hyperborea,
it is best distinguished from them by its short, scrotiform spur (though the specimen analysed by us
was the relatively slender-spurred ‗gracilis‘ morph).
Platanthera dilatata is shown as being only subtly molecularly distinct from P. viridiflora (Fig. 2),
separated by a single base pair change plus two single base pair indels. The group was considered by
both Luer (1975) and Sheviak (2002) to contain three varieties, differentiated by contrasts in the
amount of green pigment in their flowers, overall flower size and the relative dimensions of the
floral organs, notably the spur and the gynostemium. Sheviak (2002) speculated that these varieties
show a significant degree of pollinator specialization. Platanthera dilatata var. dilatata is the most
widespread taxon, stretching from Siberia across North America and southward to California and
Utah, whereas var. leucostachys occurs only in the western half of the distribution of the nominate
race, and var. albiflora has an even more restricted distribution across the Pacific northwest. The
group was represented only by var. leucostachys in the analysis of Hapeman and Inoue (1997), but here
it is represented by five accessions; of these, two (both of which were tentative identifications at the
varietal level) were collected by us and three were downloaded from GenBank as randomly selected
exemplars of 12 sequences deposited by Wallace (2002). No two of these accessions were separated
by more than 4 bp differences (i.e. levels of divergence similar to that separating P. viridiflora from P.
dilatata s.l.), but equally no two accessions were identical; all exhibited at least one sequence
difference (Fig. 2). The modest phylogenetic structure revealed links together the two samples of
var. leucostachys but not the two samples of var. dilatata s.s., one of which clusters with var. albiflora
whereas the other appears to have the potentially ancestral ITS allele (Fig. 2). Both the molecular
and morphological data suggest considerable complexity of relationships, consistent with the
hybridization and allopolyploidization events recently reported in the group (e.g. Wallace, 2002,
2003, 2006). Understanding of this taxonomic section has improved greatly between the treatments
70
of Luer (1975), Hapeman and Inoue (1997) and Sheviak (2002), a trend that will probably continue
during the next decade (R. Lauri, unpubl. res.; L. Wallace, unpubl. res.).
Sections Lacera and Tulotis

Members of the distinctive section Lacera, recognized only relatively recently by Hapeman
and Inoue (1997), have a fringed/toothed, three-lobed labellum and, in many cases, brightly
coloured flowers (the first and third features are convergent with species in the well supported
section Blephariglottis: Hapeman and Inoue, 1997). Our analysis included only three species of section
Lacera. At this level of species sampling, the tree strongly supports P. grandiflora (northern
Appalachians) as sister to P. peramoena (mid- to southern Appalachians) plus P. praeclara (US
Midwest), a topology congruent with that obtained by Hapeman and Inoue (1997). Above section
Lacera we have only the Oriental P. sonoharai as the placeholder for the widespread section Tulotis (cf.
Hapeman and Inoue, 1997).
Sections Lysiella and Lysias

As discussed at the beginning of this section, we maintain reservations regarding the
reliability of the sister relationships shown in Fig. 1 of section Lysiella and section Lysias to each
other, and when combined as sister to section Piperia. Certainly, this clade would encompass an
exceptionally broad range of morphologies, though it would permit the likely origin of section Piperia
within North America. Support for both the clade consisting of sections Lysiella, Lysias and Piperia
and the sister clade of section Platanthera is sufficiently weak that inclusion of any of the former
sections in the latter cannot confidently be refuted.
Section Piperia
Piperia was previously widely recognized as a separate genus in North America (Luer, 1975;
Wood and Neiland, 2001c; Ackerman and Morgan, 2002). However, although the ITS study of
Bateman et al. (2003) clearly demonstrated that it was strongly supported as monophyletic, it showed
equally convincingly that the clade was nested well within the genus Platanthera (see also Lauri, 2007).
Piperia was therefore relegated to a section of the genus Platanthera. The molecular branches
subtending section Piperia and the clade that it forms with sections Lysiella and Lysias are no longer
than those subtending other sections of Platanthera that were previously frequently recognized as
separate genera (Limnorchis, Lacera, and Tulotis). However, in the genus Piperia into the genus
Platanthera has met greater resistance from orchid taxonomists, primarily because Piperia is delimited
by morphological synapomorphies (notably short, inconspicuous caudicles and globose rather than
fusiform tubers: cf. Luer, 1975; Ackermann, 1977, 2002; Wood and Neiland, 2001c; Ackerman and
Morgan, 2002; Bateman et al., 2003), whereas the genus Platanthera, as presently delimited, is not.
However, we note that inconspicuous caudicles are a feature of several other genera in subtribe
Orchidinae, and globose tubers are the plesiomorphic condition in the subtribe. Rather, it is the
fusiform tubers of Pseudorchis and Platanthera, and their radical reduction in the Galearis clade, that
constitute the most convincing synapomorphies (Box et al., 2008; Efimov et al., 2009). The globose
tubers of section Piperia are ostensibly a reversion to the plesiomorphic condition within Orchidinae,
though we suspect that careful ontogenetic studies would reveal developmental differences between
the globose tubers in section Piperia vs. those of more distant genera such as Orchis s.s. We also note
that field recognition (a critical element of any successful classification) is dependent primarily on
finding reliable differences between species in one or more character states; for identification
purposes, it is unimportant whether these constitute synapomorphies.
71
Levels of sequence divergence between the three Piperia accessions included in Fig. 1 are
consistent with their status as separate species. The history of species recognition in the section
post-Rydberg (1901a, b) was summarized by Bateman et al. (2003, p. 21). However, the taxonomic
revision given by Bateman in Table 4 of Bateman et al. (2003) requires further revision, as he failed
to consider the possibility of accidentally creating orthographic synonyms when combining species
and subspecies of ‗Piperia‘ into Platanthera (J. Zarucchi, pers. comm., 2003). These errors are here
corrected in Table 1b. A current in-depth study by Lauri (2007) should provide a greatly improved
understanding of the origin and evolution of section Piperia, as well as a much needed test of
previous hypotheses that the ten constituent species of the group originated relatively recently
(Ackerman, 1977).
TABLE 1. [partial information] Taxonomic revisions of the former genera Amerorchis, [Aceratorchis,
Chondradenia, Diphylax and Tsaiorchis], plus a nomenclatural update on the former genus Piperia
(a) Former genera Amerorchis, [Aceratorchis and Chondradenia]

Galearis rotundifolia (Banks ex Pursh) R.M. Bateman, [comb. nov.]
Basionym: Orchis rotundifolia Banks ex Pursh, Fl. Amer. Sept., 2: 588 (1814).
Synonyms: Habenaria rotundifolia (Banks) Richardson, In Franklin: Journey: 750 (1823); Platanthera
rotundifolia (Banks) Lindl., Gen. Sp. Orchid.: 292(1835); Amerorchis rotundifolia Hultén, Ark. Bot. ser. 2,
7: 34 (1968).
(b) Former genus Piperia (supplement to revision by Bateman in Bateman et al., 2003)
Platanthera unalascensis (Spreng.) Kurtz, Bot. Jahrb. 19: 408 (1894).
Basionym: Spiranthes unalascensis Spreng., Systema Vegetabilium 3: 708 (1826).
Synonyms: Herminium unalasc(hk)ense (Spreng.) Rchb.f., Icon. Fl. Germ. Helv. 13–14: 107 (1838);
Platanthera foetida Geyer ex Hook.f., J. Bot. Kew Misc. 7: 376 (1855); Habenaria unalasc(h)ensis (Spreng.)
S.Wats., Proc. Amer. Acad. Arts 12: 277 (1877); Piperia unalasc(h)ensis (Spreng.) Rydberg, Bull. Torrey
Bot. Club 28: 270 (1901a).
Platanthera ephemerantha* R.M. Bateman, [nom. nov.]
Synonyms: Piperia candida R. Morgan & Ackerman, Lindleyana 5: 207 (1990); Platanthera candida (R.
Morgan & Ackerman) R.M. Bateman, nom. illegit., Bot. J. Linn. Soc. 142: 21 (2003) (illegitimate
homonym of Platanthera candida Lindl., Gen. Spec. Orchid. Pl.: 295 (1835).
Etymology: Gr. ephemera (transient) and Gr. anthus (flower): ‗The flowers in Piperia candida are
more completely white and more ephemeral than in any other member of the genus‘ (Ackerman,
2002, p. 574). The holotype of ‗Piperia‘ candida is retained, following Article 72.1a of the ICBN.
72
Die Gattungen Platanthera L.C. RICH. und Blephariglottis RAF.

in den USA und Kanada
The Genus Platanthera L.C. RICH. and Blephariglottis RAF.
in the USA and Canada
Dr. Norbert Baumbach, Michaelisstrasse 28, 99084 Erfurt
Emil Lückel, Bornemannstrasse 2, 60599 Frankfurt/Main
Originally published in Die Orchidee 60(1/2): 119-127. 2009
Reprinted by permission of senior author.
Ed. Note:
The following is a liberal translation of the text for this paper. Any errors in meaning or concept are mine. The original
paper is illustrated with beautiful full color photographs taken primarily by our own Scott Shriver of Pittsburgh, Penn.
PMB
Summary:
We propose to resurrect the North American genus Blephariglottis Raf. for the conspicuous ‗fringed Platantheras
and Habenarias‘, as already suggested by Senghas and Schrenk: genera have been kept separate for less significant features
In the literature the type Blephariglottis albiflora has been treated as a misinterpreted synonym, thinking
presumably that the varietal epithet of Orchis ciliaris var. alba, cited by Rafinesque should have been adapted. But it does
not have priority outside of its rank. He also cited Willdenow's specific epithet Blephariglottis, but this would produce an
autonym.
We think that Blephariglottis longicornis Raf. - by its Latin description and given range - agrees with the integrilabia
concept; but realize that the less typical lip may warrant a different genus placement.
In the North American orchid flora, there are a number colourful and handsome species that 20/21 st century
American authors place in the genus Platanthera L.C. Rich., although they differ significantly in generic concepts. The
resulting group, recognized native within the genus Platanthera in the United States and Canada now includes 38 species
and 11 natural hybrids (P.M. Brown 2003). [I am not sure how the authors arrived at 38 species; currently (2009) there
are 34 species, including several new ones, and 21 hybrids in Platanthera s.l. Ed.]
The genus Platanthera was created in 1818 by L.C. Richard. The type of the genus is the
Eurasian Platanthera bifolia (L.) L.C. Rich. which is not known from North America. The original
description indicated characteristics such as are the pair of opposite basal leaves and often narrow
one-lobed lip with spur.
Willdenow 1805 and J. Lindley (1835) transferred a number of species from the genera Orchis
(L) and Habenaria to the new genus Platanthera. C.S. Rafinesque created in 1836 (publ. 1837) the
genus Blephariglottis with a total of five species from the United States and the type species
Blephariglottis albiflora Raf. Rydberg (1901) and Farwell (1901) added more species. Characteristic
features of the genus are the partly three lobed lip, which is ciliated or toothed (sometimes only a
little bit visible), and the stem with several leaves. Striking features are the large, usually intensely
coloured flowers.
Until the first third of the last century, the genus Blephariglottis was recognized, then was
merged with Platanthera and Blephariglottis into the compilogenus genus Habenaria by the American
modern taxonomists Ames, Correll and Schweinfurth. Only Luer (1972, 1975) separated the genus
Platanthera from Habenaria but treated Blephariglottis as a section, which until today is the view taken in
North American literature.
73
Schrenk (1976, 1977a) and Senghas (1973) presented a carefully selected 'balanced proposal
for the allocation of multiple genera within Habenaria’, and included in the consideration was the
genus Blephariglottis. Blephariglottis alba is considered in the literature as an incorrect species name
because the epithet "alba" is a basionym of a variety and not a species, so Rafinesque created a
legitimate new epithet be used. Consequently, Blephariglottis albiflora Raf. is the valid name.
Of the five species which Rafinesque described, four species were later recognized as such.
One taxon, however, Blephariglottis longicornis Raf. remained overlooked in all subsequent publications.
The only reference takes place in the World Checklist (Kew) Govaerts, where the species is listed as
a synonym for the Indian Habenaria longicornis Lindl. 1835, which is not accurate because it is a
completely different species.
The Latin description of Rafinesque is "Elatior, fol. obtusiusc. nervosis, spica oblonga, petalis albis, 2
internis apic ciliatis, calcar longissimo divaricato, labello lineari, basi ciliis brevis, apic lacero fimbriato - Alabama,
Louisiana." According to this description and the distribution of this species is clearly Blephariglottis
integrilabia (Correll) Schrenk 1977a which will become synonymous with this taxon because of the
nomenclature rules [There is no documentation or report for Louisiana other than Rafinesque‘s description and this
may be the basis for not embracing his earlier name. Ed.]
The three-parted lipped species of this group were placed in the genus Fimbriella Farw. ex
Butzin by Butzin (1981) – with the type Fimbriella lacera (Michx.) Butzin. After DNA analysis by
Bateman et al. (1997) and Pridgeon et al. (1997), results for the genus Platanthera previously had gaps
and analysis of only a few species had been made, and the result is the type species which is directly
related to the three-parted lipped species from Asia (e.g. Platanthera florentia) and the North American
species. For groups of other species i.e. Blephariglottis ciliaris (Platanthera ciliaris) as well as for the
Platanthera group and Limnorchis no conclusive statements have been made, so currently the
resurrection of the genus Blephariglottis based upon morphology appears justified.
New combinations for formae are proposed here:

Blephariglottis albiflora Raf. forma holopetala (Lindl.) Baumbach & P.M. Brown comb.
nov.
Basionym: Platanthera holopetala Lindl. Gen. Sp. Orchid. Pl. 291. 1835.
Blephariglottis cristata Raf. forma straminea (P.M. Brown) Baumbach & P.M. Brown comb.
nov.
Basionym: Platanthera cristata Lindl. forma straminea P.M. Brown North Amer. Native Orchid J., 1(1): 12. 1995.
Blephariglottis grandiflora (Lindley) Rydb. forma albiflora (Rand & Redf. ) Baumbach &
P.M. Brown comb. nov.
Basionym: Habenaria fimbriata (Aiton) R. Br. forma albiflora Rand. & Redf. Mt. Desert Isl. 153. 1894.
Blephariglottis grandiflora (Bigelow) Rydb. forma bicolor (P.M. Brown) Baumbach & P.M.
Brown comb. nov.
Basionym: Platanthera grandiflora forma bicolor P.M. Brown N. Amer. Native Orchid J. 1(1): 12. 1995.
Blephariglottis grandiflora (Lindl.) Rydb. forma carnea (P.M. Brown) Baumbach & P.M.
Brown comb. nov.
Basionym: Platanthera grandiflora forma carnea P.M. Brown N. Amer. Native Orchid J. 1(1): 12. 1995.
Blephariglottis grandiflora (Bigelow) Rydb. forma mentotonsa (Fernald ) Baumbach & P.M.
Brown comb. nov.
Basionym: Habenaria fimbriata R. Br. forma mentotonsa Fernald Rhodora 48: 184. 1946.
74
Blephariglottis peramoena (A. Gray) Rydb. forma doddsiae (P.M. Brown) Baumbach &
Basionym: Platanthera peramoena (A. Gray) A. Gray forma doddsiae P.M. Brown N. Amer. Native Orchid J. 8: 30 (71, t. 3
(bottom)). 2002.
Blephariglottis psycodes (L.) Rydb. forma albiflora (R. Hoff.) Baumbach & P.M. Brown
comb. nov.
Basionym: Habenaria psycodes (L.) Spreng. forma albiflora R. Hoffm. Proc. Boston Soc. Nat. Hist. 36: 248. 1922.
Blephariglottis psycodes (L.) Rydb. forma fernaldii (J. Rousseau & Rouleau) Baumbach &
Basionym: Habenaria psycodes (L.) Spreng. var. fernaldii J. Rousseau & Rouleau. Bull. Jard. Bot. État Bruxelles 27: 370.
1957.
Blephariglottis psycodes (L.) Rydb. forma ecalcarata (Bryan) Baumbach & P.M. Brown
comb. nov.
Basionym: Habenaria psycodes (L.) Spreng. var. ecalcarata Bryan Ann. Missouri Bot. Gard. 4: 38. 1917.
Blephariglottis psycodes (L.) Rydb. forma rosea (P.M. Brown) Baumbach & P.M. Brown
comb. nov.
Basionym: Platanthera psycodes (L.) Lindl. forma rosea P.M. Brown North Amer. Native Orchid J., 1(4): 289. 1995.
Blephariglottis psycodes (L.) Rydb. forma varians (Bryan) Baumbach & P.M. Brown comb.
nov.
Basionym: Habenaria psycodes (L.) Spreng. var. varians Bryan Ann. Missouri Bot. Gard. 4: 38. 1917.
The genus Blephariglottis in North America as treated by Baumbach and Lückel.

Blephariglottis albiflora Raf. forma carnea (P.M. Brown) Baumbach & P.M.
Syns: Platanthera blephariglottis (Willd.) Lindl., Habenaria Brown
blephariglottis (Willd.) Hooker forma mentotonsa (Fernald ) Baumbach & P.M.
forma holopetala (Lindl.) Baumbach & P.M. Brown Brown
Blephariglottis chapmanii Small Blephariglottis longicornis Raf. Syns.: Platanthera
Syns: Platanthera chapmanii (Small) Luer, integrilabia (Correll) Luer, Habenaria blephariglottis var.
Habenaria xchapmanii (Small) Ames integrilabia Correll, Habenaria correlliana Cronquist
Blephariglottis ciliaris (L.) Rydb. Blephariglottis lacera (Michx.) Farw.
Syns.: Platanthera ciliaris Lindl., Habenaria ciliaris Syns.: Platanthera lacera (Michx.) G. Don, Habenaria lacera
(L.) R. Brown Michx. R. Brown
Blephariglottis conspicua (Nash) Small Blephariglottis leucophaea (Nutt.) Farw.
Syns.: Platanthera conspicua (Lindl.) P.M. Brown, Syns.: Platanthera leucophaea (Nutt.) Lindl., Habenaria
Habenaria conspicua Nash (not treated in Baumbach & leucophaea (Nutt.) Gray
Lückel) Blephariglottis pallida (P.M. Brown) Baumb. &
Blephariglottis cristata Raf. Lueckel
Syns.: Platanthera cristata Michx., Syn.: Platanthera pallida P.M. Brown
Habenaria cristata (Michx.) R. Brown Blephariglottis peramoena (A. Gray) Rydb.
forma straminea (P.M. Brown) Baumbach & P.M. Syns.: Platanthera peramoena (A. Gray) A. Gray, Habenaria
Brown peramoena A. Gray
Blephariglottis grandiflora (Bigelow) Rydb. forma doddsiae (P.M. Brown) Baumbach & P.M.
Syns.: Platanthera grandiflora (Bigelow) Lind., Habenaria Brown
grandiflora (Bigelow) Torrey ex L.C. Beck Blephariglottis praeclara (Sheviak & Bowles)
forma albiflora (Rand & Redf. ) Baumbach & P.M.
Baumb. & Lueckel
Brown Syns.: Platanthera praeclara Sheviak & Bowles, Habenaria
forma bicolor (P.M. Brown) Baumbach & P.M. leucophaea A. Gray var. praeclara (Sheviak & Bowles)
Brown Cronquist
75
Blephariglottis psycodes (L.) Rydb. Blephariglottis ×canbyi (Ames) W. Stone

Syns.: Platanthera psycodes (L.) Lindl., Habenaria psycodes Syns.: Platanthera ×canbyi (Ames) Luer, Habenaria ×canbyi
(L.) Sprengl. Ames
forma albiflora (R. Hoff.) Baumbach & P.M. Blephariglottis ×channellii (Folsom) Baumb. &
Brown Lueckel
forma fernaldii (J. Rousseau & Rouleau) Baumbach Syn.: Platanthera ×channellii Folsom
& P.M. Brown Blephariglottis ×enigma (P.M. Brown) Baumb. &
forma ecalcarata (Bryan) Baumbach & P.M. Brown Lueckel
forma rosea (P.M. Brown) Baumbach & P.M. Syn.: Platanthera ×enigma P.M. Brown
Brown Blephariglottis ×hollandiae (Catling & Brownell)
forma varians (Bryan) Baumbach & P.M. Brown Baumb. & Lueckel
Blephariglottis shriveri (P.M. Brown) Baumb. & Syn.: Platanthera ×hollandiae Catling & Brownell
Lueckel Blephariglottis ×keenanii (P.M. Brown) Baumb.
Syn.: Platanthera shriveri P.M. Brown & Lueckel
Blephariglottis ×andrewsii (M. White) House, Syn.: Platanthera ×keenanii P.M. Brown
Syns.: Platanthera ×andrewsii (M. White) Luer, Habenaria Blephariglottis ×lueri (P.M. Brown) Baumb. &
×andrewsii M. White
Blephariglottis ×apalachicola (P.M. Brown & S.
Lueckel
Syn.: Platanthera ×lueri P.M. Brown
Stewart) Baumb. & Lueckel
Syn.: Platanthera ×apalachicola P.M. Brown & S. Stewart Blephariglottis ×osceola (P.M. Brown & S.
Blephariglottis ×beckneri (P.M. Brown) Baumb. Stewart) Baumb. & Lueckel
Syn.: Platanthera ×osceola P.M. Brown & S. Stewart
& Lueckel
Syn.: Platanthera ×beckneri P.M. Brown Blephariglottis ×reznicekii (Catling, Brownell &
Blephariglottis ×bicolor Raf. Allen) Baumb. & Lueckel
Syn.: Platanthera ×bicolor (Raf.)Luer Syn.: Platanthera ×reznicekii Catling, Brownell & Allen
SEED MICROMORPHOLOGY SUPPORTS THE SPLITTING OF

LIMNORCHIS FROM PLATANTHERA (ORCHIDACEAE)
Roberto Gamarra, Pablo Galán, Irene Herrera and Emma Ortúñez
Originally published in Nordic Journal of Botany 26: 61-65, 2008
Reprinted by permission of publisher.
Seeds of several species of the genera Platanthera and Habenaria have been studied by means of scanning
electron microscopy. In the genus Platanthera, two morphological patterns were found. One of these appears in the so-
called ‗dilatata-hyperborea complex‘. It is proposed that this pattern is a good diagnostic character to split off this group of
species into the genus Limnorchis. In Habenaria, the morphological patterns vary between the species and differs
completely from those found in Platanthera.
According to Pridgeon et al. (2001), the genus Platanthera (L.) Rich. comprises about 200
species, and the genera Tulotis Raf., Blephariglottis Raf., Limnorchis Rydb., Lysiella Rydb., Gymnadeniopsis
Rydb. and Piperia Rydb. are included as synonyms. It belongs to the subtribe Orchidinae (Dressler
1993). Platanthera is distributed throughout the north temperate area, extending their distribution to
Central America, northern Africa and New Guinea. Most of the species occur in North America and
Asia. Habenaria Willd. comprises about 600 species in tropical and temperate regions, principally in
America and Africa (Buttler 2001). It belongs to the subtribe Habenariinae (Dressler 1993). Several
North American taxa such as Platanthera dilatata (Pursh) Lindl. and P. hyperborea (L.) Lindl. are
76
included in the ‗dilatata-hyperborea‘ complex. This critical group was assigned to the genus Limnorchis
by Rydberg (1900, 1901) and this taxonomical treatment was followed by several authors like Britton
and Brown (1943) and Löve and Simon (1968). However, Ames (1910), Mohlenbrock (1970) and
Voss (1972) included the species of this group in the genus Habenaria. On the other hand, Luer
(1975) recognised this complex as the section Limnorchis within the genus Platanthera and this view
was later accepted by Hapeman and Inoue (1997) and Sheviak (1999a). Sheviak (1999b) proposed
the new name P. aquilonis for the North American populations previously referred to as P. hyperborea,
restricting the usage of the latter name to a species that only occurs in Greenland and Iceland.
Therefore, the actualised name of this complex must be the ‗dilatata-aquilonis complex‘. The former
inclusion by American botanists (Ames 1910, Correll 1950) of North American species of Platanthera
in the genus Habenaria is misapplied. All contemporary specialists on the Orchidaceae recognize
Platanthera and Habenaria as distinct genera (Luer 1975, Case 1987, Smith 1993) and, in fact, the
genus Habenaria is referred to the subtribe Habenariinae whereas Platanthera belongs in subtribe
Orchidinae (Dressler 1993, Pridgeon et al. 2001).
Other North American species of the genus Platanthera, such as P. unalascensis (Spreng.) Kurtz
and P. elegans Lindl. were included in the genus Piperia by Rydberg (1901), and Luer (1975) and
Ackermann (1977) recognised three and four species each in this genus, respectively. The last author
distinguished Piperia from Platanthera by several morphological characters (globose tubers vs.
fusiform tubers, very short and inconspicuous caudicles, senescence of leaves during anthesis).
However, in Europe no segregate genera have been proposed and the genus Platanthera has been
traditionally circumscribed (Webb 1980, Delforge 1994).
In the phylogenetic analysis of Hapeman and Inoue (1997), using the ITS regions, the genus
Platanthera is monophyletic and distant from Habenaria s. s. In this analysis, five major clades could
be recognized within Platanthera and one of them agrees with the section Limnorchis. However,
according to these authors, the relationships between the clades are weakly supported.
Simultaneously, Bateman et al. (1997), obtained a different topology that indicates that the
genus is polyphyletic, with P. hyperborea being the most divergent of the studied species, although the
studied material of this taxon is from North America and thus rather belongs to P. aquilonis. Later,
Bateman et al. (2003) proposed to encompass the genera Piperia and Limnorchis within Platanthera.
Healey et al. (1980) observed two distinct seed morphological patterns for the studied
species of Platanthera and Piperia. The seeds of the genus Piperia agree with the samples studied by us
in the European species of Platanthera. However, the three species of Platanthera (P. dilatata, P.
hyperborea, P. saccata) studied by these authors, all of which belong in the ‗dilatata-aquilonis complex‘,
have smooth seeds, as also shown in the present study.
During our research about seed micromorphology of the European genera in the subtribe
Orchidinae, we have observed a strong agreement between the qualitative characters and the genera
(Gamarra et al. 2007). According to this criterion, Platanthera has reticulate seeds with a truncate
apical zone but without lamella. This pattern is also found within the species previously referred to
Piperia.
Our results largely support the result of the molecular phylogeny published by Bateman et al.
(2003), who nested the genus Piperia within Platanthera. On the contrary, the studied species of the
77
‗dilatata-aquilonis complex‘ have smooth seeds with a rounded apical zone and presence of lamella.
This pattern supports splitting off this group from Platanthera to include it in the segregate genus
Limnorchis, and these results are consistent with the paraphyletic Platanthera obtained by Bateman et
al. (1997) and with the separate clade formed by Limnorchis in the analysis by Hapeman and Inoue
(1997). The testa features observed by us in species of Platanthera s. l. (including Limnorchis) are
completely different from those found in the genus Habenaria, which show a great intrageneric
variability. This result supports the polyphyletic character of the genus Habenaria revealed by
Bateman et al. (2003).
Ed.: No new combinations were made in the above paper. Those combinations needed are
proposed below:
Limnorchis aquilonis (Sheviak) Rebrist. & Elven forma alba (Light) P.M. Brown, S.L.
Stewart & R. Gamarra comb. nov.
Basionym: Platanthera hyperborea (L.) Lindley forma alba M.H.S. Light Lindleyana 4: 158 (-160; figs. 1-3). 1989.
Syn.: Platanthera aquilonis Sheviak alba (Light) P.M. Brown
Limnorchis brevifolia (Greene) Rydb. forma alba (P.M. Brown) P.M. Brown, S.L. Stewart &
R. Gamarra comb. nov.
Basionym: Platanthera brevifolia (Greene) Kraenzl. forma alba P.M. Brown McAllen Int. Orchid Soc. J. 7(12): 8 (-9; fig. 3).
2006.
Limnorchis dilatata (Pursh) Rydb. var. leucostachys (Lindl.) P.M. Brown, S.L. Stewart & R.
Gamarra comb. nov.
Basionym: Limnorchis leucostachys (Lindl.) Rydb. Mem. New York Bot. Gard. 1: 106. 1900.
Syns.: Platanthera dilatata (Pursh) Lindl. var. leucostachys (Lindl.) Luer, Habenaria dilatata var. leucostachys (Lindl.) Ames
Limnorchis holochila (Hillebr.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym. Habenaria holochila Hillebr. Fl. Haw. Isl. 432. 1888.
Syn.: Platanthera holochila (Hillebr.) Kraenzl. (known only from Hawaii)
Limnorchis limosa (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera limosa Lindl. Ann. Nat. Hist. 4(26): 381. 1840.
Syn.: Habenaria limosa (Lindl.) Hemsl.
Limnorchis tescamnis (Sheviak & W.F. Jenn.) P.M. Brown, S.L. Stewart & R. Gamarra comb.
nov.
Basionym: Platanthera tescamnis Sheviak & W.F. Jenn. Rhodora 108(933): 20 (19-31; figs. 1-2). 2006.
Limnorchis tipuloides (L. f.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Orchis tipuloides L. f. Suppl. Pl. 401. 1782 [1781 publ. Apr 1782]
Syns.: Platanthera tipuloides (L. f. ) Lindl., Habenaria tipuloides (L. f. ) Bentham
Limnorchis tipuloides ( L. f.) P.M. Brown, S.L. Stewart & R. Gamarra var. behringiana
(Rydb.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Limnorchis behringiana Rydb. Bull. Torrey Bot. Club 1901, 620
Syn.: Platanthera tipuloides (L. f.) Lindl. var. behringiana (Rydb.) Hultén
78
Limnorchis yosemitensis (Colwell, Sheviak & P.E. Moore) P.M. Brown, S.L. Stewart & R.
Gamarra comb. nov.
Basionym: Platanthera yosemitensis Colwell, Sheviak & P.E. Moore Madroño 54(1): 86 (-93; figs. 1-2). 2007.
Limnorchis ×correllii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×correllii W.J. Schrenck Die Orchidee (Hamburg) 26(6): 262. 1975.
Limnorchis ×estesii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×estesii W.J. Schrenck Die Orchidee (Hamburg) 26(6): 262. 1975.
Limnorchis ×evansiana (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×evansiana P.M. Brown North American Native Orchid Journal 15(1): 40. 2009.
Limnorchis ×folsomii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×folsomii P.M. Brown North American Native Orchid Journal 15(1): 40. 2009.
Limnorchis ×kelleyi (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×kelleyi P.M. Brown North American Native Orchid Journal 15(1): 40. 2009.
Limnorchis ×lassennii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×lassenii W.J. Schrenk Die Orchidee (Hamburg) 26(6): 262. 1975.
Limnorchis ×smithii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Basionym: Platanthera ×smithii P.M. Brown North American Native Orchid Journal 15(1): 40. 2009.
The Genus Limnorchis in the United States and Canada.

Synonyms for those species treated within Platanthera and Habenaria are also given.
Limnorchis aquilonis (Sheviak) Rebrist. & Elven Limnorchis dilatata (Pursh) Rydb. var. albiflora
syn.: Platanthera aquilonis Sheviak (Cham.) Efimov
forma alba (Light) P.M. Brown, S.L. Stewart & R. Syns.: Platanthera dilatata (Pursh) Lindl. var. albiflora
Gamarra (Cham.) Ledeb., Habenaria dilatata var. albiflora (Cham.)
Correll
Limnorchis brevifolia (Greene) Rydb. Limnorchis dilatata (Pursh) Rydb. var. leucostachys
Syns.: Platanthera brevifolia (Greene) Kranzl., Habenaria (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra
brevifolia Greene Syns.: Limnorchis leucostachys (Lindl.) Rydb., Platanthera
forma alba (P.M. Brown) P.M. Brown, S.L. Stewart dilatata (Pursh) Lindl. var. leucostachys (Lindl.) Luer,
& R. Gamarra Habenaria dilatata var. leucostachys (Lindl.) Ames
Limnorchis chorisiana (Cham.) J.P. Anderson
Syns. Platanthera chorisiana (Cham.) Reich. f., Habenaria Limnorchis huronensis (Nutt.) Rydbg.
chorisiana Cham. Syns.: Platanthera huronensis (Nutt.) Lindl., Habenaria
huronensis (Nutt.) Spreng.
Limnorchis convallariifolia (Fischer ex Lindl.) Rydb.
Syns.: Platanthera convallariifolia Fischer ex Lindl., Habenaria Limnorchis hyperborea (L.) Rydb.
convallariifolia (Lindl.) B. Boivin Syns.: Platanthera hyperborea (L.) Lindl., Habenaria hyperborea
(L.) R. Br. ex W.T. Aiton (known only from Greenland
Limnorchis dilatata (Pursh) Rydb. var. dilatata and Iceland)
Syns.: Platanthera dilatata (Pursh) Lindl. var. dilatata,
Habenaria dilatata (Pursh) Hooker
79
Limnorchis holochila (Hillebr.) P.M. Brown, S.L. Limnorchis viridiflora ( Cham. ) Rydb.
Stewart & R. Gamarra Syns: Platanthera hyperborea var. viridiflora (Cham.) Luer,
Syns.: Habenaria holochila Hillebr., Platanthera holochila Habenaria borealis Cham. var. viridiflora Cham. (known
(Hillebr.) Kraenzl. (known only from Hawaii) from western Alaska per Bateman et al.)
Limnorchis yosemitensis (Colwell, Sheviak & P.E.
Limnorchis limosa (Lindl.) P.M. Brown, S.L. Moore) P.M. Brown, S.L. Stewart & R. Gamarra
Stewart & R. Gamarra Syn.: Platanthera yosemitensis Colwell, Sheviak & P.E.
Syns.: Platanthera limosa Lindl., Habenaria limosa (Lindl.) Moore
Hemsl.
Limnorchis zothecina (L.C. Higgins & S.L. Welsh)
Limnorchis purpurascens Rydb. W.A. Weber
Syns.: Platanthera purpurascens (Rydb.) Sheviak & W.F. Syns.: Platanthera zothecina (L.C. Higgins & S.L. Welsh)
Jenn., Habenaria purpurascens (Rydb.) Tidestr. in Tidestr. & Kartesz & Gandhi, Habenaria zothecina L.C. Higgins & S.L.
Kittell Welsh
Limnorchis sparsiflora (S. Watson) Rydb. Limnorchis ×correllii (W.J. Schrenck) P.M. Brown,
Syns.: Platanthera sparsiflora (S. Watson) Schltr., Habenaria S.L. Stewart & R. Gamarra
sparsiflora S. Watson Syn.: Platanthera ×correllii W.J. Schrenck
Limnorchis ×estesii (W.J. Schrenck) P.M. Brown,
Limnorchis stricta (Lindl.) Rydb. S.L. Stewart & R. Gamarra
Syns.: Platanthera stricta Lindl., Habenaria stricta (Lindl.) Syn.: Platanthera ×estesii W.J. Schrenck
Rydb.
Limnorchis ×evansiana (P.M. Brown) P.M. Brown,
S.L. Stewart & R. Gamarra
Limnorchis tescamnis (Sheviak & W.F. Jenn.) P.M. Syn.: Platanthera ×evansiana P.M. Brown
Brown, S.L. Stewart & R. Gamarra Limnorchis ×folsomii (P.M. Brown) P.M. Brown,
Syn.: Platanthera tescamnis Sheviak & W.F. Jenn.
Syn.: Platanthera ×folsomii P.M. Brown
Limnorchis tipuloides (L. f.) P.M. Brown, S.L. Limnorchis ×kelleyi (P.M. Brown) P.M. Brown,
Stewart & R. Gamarra S.L. Stewart & R. Gamarra
Syns: Platanthera tipuloides (L. f. ) Lindl., Habenaria tipuloides Syn.: Platanthera ×kelleyi P.M. Brown
(L. f. ) Bentham
Limnorchis ×lassennii (W.J. Schrenck) P.M.
Limnorchis tipuloides ( L. f.) P.M. Brown, S.L. Brown, S.L. Stewart & R. Gamarra
Stewart & R. Gamarra var. behringiana (Rydb.) Syn.: Platanthera ×lassenii W.J. Schrenk
P.M. Brown, S.L. Stewart & R. Gamarra Limnorchis ×smithii (P.M. Brown) P.M. Brown,
Syns.: Limnorchis behringiana Rydb., Platanthera tipuloides (L.
f.) Lindl. var. behringiana (Rydb.) Hultén
Syn.: Platanthera ×smithii P.M. Brown
What does all of this mean?

 If you embrace the generic concept of Blephariglottis you must use the species epithets set
forth for that genus with significant changes being B. albiformis for P. blephariglottis and B.
longicornis for P. integrilabia and the new combinations completing that genus. Be aware that
there are no published combinations for the species epithets albiformis or longicornis in the
genera Habenaria or Platanthera. If you disagree with the use of the two new species epithets
and still wish to use the genus Blephariglottis you may use B. integrilabia (Correll) Schrenk but
must still use B. albiformis for reasons set forth earlier in the above paper.
 If you embrace the generic concept of Limnorchis you must use the combinations set forth
for that genus with no significant changes in specific epithets, but using new combinations
completing that genus.
80
 If you embrace the concept of merging of Amerorchis into Galearis you must use the new
combination Galearis rotundifolia.
 If you embrace the concept of merging Piperia into Platanthera you must use the new
combinations and names set forth for that genus with a significant change for Piperia candida
to Platanthera ephemerantha. One of the more interesting concepts in Platanthera is the
possibility of recognizing P. viridiflora as a species. Not in the concept of a synonym for P.
stricta sensu Sheviak but as a far northwestern (western Alaska) counterpart(?) species to
extreme northeastern P. hyperborea (Greenland and Iceland).
ACKNOWLEDGEMENTS:
Annals of Botany: Dr. David Frost
Nordic Journal of Botany: Dr. Maria Persson, Dr. Roberto Gamarra
Die Orchidee: Dr. Norbert Baumbach, Dietrich and Ursula Rückbrodt for assistance in the translation
Glossary assistance from Lucy Dueck and Jim Fowler
Photo of Limnorchis holochila by Dr. Lawrence W. Zettler
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Native Orchid Journal 14(4):255-261
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83
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Luer, C. A. (1975): The native Orchids of the United States and Canada Excluding Florida :75-93: 145-148
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GLOSSARY
allele: one of two or more alternative forms of a gene, regions and its length is not a multiple of 3, an
occupying the same position (known as a indel will cause a frameshift mutation,
locus) on paired chromosomes and possibly resulting in the production of a
controlling the same inherited characteristics; different protein
alternative DNA sequences at the same locus Internal Transcribed Spacer (ITS): refers to a piece of
which may or may not result in different non-functional RNA situated between
phenotypic traits structural ribosomal RNAs (rRNA) on a
allopolyploidy: the condition of having two or more common precursor transcript
complete sets of chromosomes derived from lamella: layers of membranous material often
different species containing polygalacturons (D-galaturonic
atavistic: relating to or displaying the recurrence of a acid) and neutral carbohydrates (pectins)
genetic feature that has been absent for monophyletic: containing all descendants and their
several generations; the tendency to revert to most recent common ancestor in one clade
an ancestral type monotypic: a genus with only one species
autogamy: self-pollinations whereby the fertilizing nominate race: the original form or race for which the
pollen is from the same blossom as is the group was named
pistil receptacle orthographic: relating to spelling or wording
basionym: the original previously published legitimate paedomorphosis: phylogenetic change that involves
name or epithet from which a new name is retention of juvenile characters by the adult ;
suggested for a taxon of different rank or also, a phenotypic and/or genetic change in
position which the adults of a species retain traits
bursicle: a small sac or pouch-like receptacle previously seen only in juveniles
clade: a group of organisms consisting of a single paraphyletic: a paraphyletic group contains some, but
common ancestor and all its descendants; a not all, of the descendants from a common
monophyletic group ancestor; containing the most recent common
gynostemium: a reproductive structure found in several ancestor but not all its descendants in one
plant families whereby the male and female clade
parts (stamens and pistil) are fused into a parsimonious: very frugal or ungenerous; the most
single organ; column conservative approach with the least number
heterochrony: changes in the timing or relative rates of of changes
development of different tissues within an phylogenetic: genetic evolutionary relationships within
organism that can be inherited by its offspring and between taxonomic levels, particularly the
which result in changes to size and shape patterns of descent, often branching, from
homoplastic: analogous/similar features or characters one organism to another
that are independently derived within separate plesiomorphic: An ancestral or primitive character
lineages which are not descended from a useful in defining a monophyletic group
recent common ancestor (e.g., octopus eyes polyphyletic: containing members whose most recent
and human eyes) common ancestor is not a member of the
indels: heritable insertions or deletions of a segment of group (e.g., warm-blooded animals containing
DNA in a sequence; can be used as a genetic mammals and birds but their last common
marker to differentiate among individuals or ancestor was not warm-blooded)
groups of individuals; if found in coding
84
section (as in section of a genus): a genetically or between the many tandemly repeated copies
morphologically distinct group of organisms of the ribosomal RNA genes
within a specified genus subtribe: a taxonomic division of a tribe
sensu lato (s.l.): in the broadest sense synapomorphies: apomorphy (derived or specialized
sensu stricto (s.s.): in the narrowest sense character) shared by two or more groups
sequence (verb): to determine the nucleotide (bases which originated in their last common
a,c,g,t) composition of a strand of DNA ancestor
usually limited in length; since DNA is synonym: a difference and equally valid scientific name
double-stranded, both strands are usually used for a single taxon
sequenced and compared to each other for testa: seed coat developed from the tissue surrounding
accuracy since each base only matches up to the ovule
one other base on the opposite strand trees: a colloquial term used to describe the
spacer (DNA): The dNA sequence between genes; the evolutionary relationships among a group of
term is used particularly for the spacer DNA organisms, much as a family tree is used to
document the genealogy of human lineages
Typical habit of
Blephariglottis (leucophaea) and Limnorchis (aquilonis)
85
Blephariglottis chapmanii Blephariglottis lacera Blephariglottis pallida

syn.: Platanthera chapmanii syn.: Platanthera lacera syn.: Platanthera pallida
Blephariglottis praeclara Blephariglottis peramoena Blephariglottis ×channellii

syn.: Platanthera praeclara syn.: Platanthera peramoena syn.: Platanthera ×channellii
syn.: Platanthera
Blephariglottis ×lueri Blephariglottis ×andrewsii

syn.: Platanthera ×lueri syn.: Platanthera ×andrewsii
86
Limnorchis chorisiana
syn.: Platanthera chorisiana
Limnorchis aquilonis
syn.: Platanthera aquilonis
Limnorchis brevifolia Limnorchis limosa

syn.: Platanthera brevifolia syn.: Platanthera limosa
Limnorchis dilatata Limnorchis yosemitensis Limnorchis holochila L. hyperborea (Greenland)

var. leucostachys syn.: Platanthera yosemitensis (Hawai‘i) syn.: Platanthera hyperborea
syn.: Platanthera dilatata var. leucostachys syn.: Platanthera holochila
87
Rydberg: LIMNORCHIS

Per Axel Rydberg
a facsimile reprint
The following is a facsimile reprint of Rydberg‘s original work on

Limnorchis from the Bulletin of the Torrey Botanical Club, Vol. 28, No.
11. (Nov., 1901), pp. 605-643 and reprint by permission. With the
increased interest in the Limnorchis as a genus many of Rydberg‘s
original observations are not only relevant but help to clarify the
position of many of these species.
It is essential to note that this was written in 1901 and much has
also changed in more than a century. Some of the species he
describes were merely aberrant specimens and others extreme
expressions of larger concepts. For the best synonymy see
Shevaik‘s treatment of Platanthera in Flora of North America vol. 26.
In this paper Rydberg enumerated 24 species of Limnorchis. 12 are
recognized today at the species level as well as 5 additional
species.
Some, but certainly not all, of the taxonomic issues that should
be noted include:
L. viridiflora was considered within L. stricta by Sheviak but a
recent publication by Bateman et al. indicates that it may be a
good species.
L. brachypetala, L. gracilis fall under L. stricta
L. ensifolia & L. laxiflora fall under L. sparsiflora
L. major, media = L. huronensis
L. arizonica = L. limosa
Much of what was identified as L. ensifolia in the Great Basin is L.
tescamnis
Much of what was identified as L. hyperborea in the continental US
and Canada is L. aquilonis
L. leptoceratitis, L. foliosa, leucostachys fall in L. dilatata and its
varieties
L. laxiflora = L. sparsiflora
88
Rydberg: LIMNORCHIS
89
Rydberg: LIMNORCHIS
90
Rydberg: LIMNORCHIS
91
Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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Rydberg: LIMNORCHIS
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116
WILD ORCHIDS IN THE 21ST CENTURY
WILD ORCHIDS IN THE 21ST CENTURY:

FIELD GUIDES AND FLORAS ON
NORTH AMERICAN NATIVE ORCHIDS
2007-2008 UPDATE
Since the publications of Wild Orchids in the 21st Century: past, present, and
future field guides and floras on North American native orchids in August 2006 (NANOJ 12:
11-28) several new works have been published. They are presented here in the same
format as the previous article.
Wild Orchids of the Northeast: New England,

New York, Pennsylvania, and New Jersey
Paul Martin Brown with original artwork by Stan Folsom
2007 University Press of Florida.
384 pages 6x9‖ field guide; 331 full color photographs, 99
line drawings, 86 maps, keys for identification;
considerable additional informational material
$29.95 Paper (Flexibind): ISBN: 0-8130-9780813030340
Field Guide to the Wild Orchids of Texas

Paul Martin Brown with original artwork by Stan Folsom
2008 University Press of Florida
316 pages 6 x 9‖ field guide; 286 full color photographs, 91
line drawings, 67 maps, keys for identification; considerable
additional informational material
$29.95 Paper (Flexibind): ISBN 978-0-8130-3159-0
117
by Chelsea Kiefer
$19.50
A young naturalist's photographic
essay recounting her discovery of
photography and orchids in
northeastern North America.
Order directly from the
publisher at
http://www.lulu.com/content/123
2040
or from the author at
chelseakieffer@gmail.com
By Sylvain Beauséjour
Preface by Marie Tifo
Foreword and English
Translation by Paul Martin Brown
Each of the 51 Quebec orchids

presented is shown by two full
pages of coloured photographs
accompanied by an explanatory
text. 176 pages 380 photographs:
11‘‘ x 9.5‘‘ Hard-bound $49.99
CAD
http://www.orchideequebec.com
118
ORQUÍDEAS DE LA REPÚBLICA
DOMINICANA Y HAITÍ
ORCHIDS OF THE DOMINICAN
REPUBLIC AND HAITI
Eladio Fernández, Francisco Jiménez, Juan
Llamacho, and James Ackerman
American Chamber of Commerce of the Dominican
Republic
209 pages, 100 illustrations
Limited printing of 2000 copies

9.25 x 12.5‖; $95 US plus shipping at cost
Ordering information: Amcham@amcham.org.do or
http://www.orchidsbooks.com/book.asp?id=1150
of interest……
Orchid Seed Germination Media,
a Compendium of Formulations
Aaron J. Hicks with Katy Lynn (editor)
The Orchid Seedbank Project
210 pages ISBN 0-9673049-2-X
$60 postpaid Order from: The Orchid Seedbank Project at
http://members.cox.net/ahicks51/osp
119
2007 2007 2008
by
Paul Martin Brown
$9.95 each
16 panels, full-color
with basic keys
The first 5 laminated

pocket guides from
the University of Iowa
Press of a projected
series of 12 guides
covering all of the
orchids of the
continental US and
Canada
Order from
naorchid@aol.com
2008 2009
120
Forthcoming
Genera Orchidacearum, Volume 5:

Epidendroideae (Part 2)
Edited by Alec M Pridgeon, Phillip J Cribb, Mark W Chase

and Finn Rasmussen
664 pages, 400 line illustrations and 48 color plates.
Oxford University Press
Hardcover due October 2009 $195
ISBN-13: 9780198507130
121
Empiricist :…..OF CABBAGES AND KINGS
…..OF CABBAGES AND KINGS

The Slow Empiricist
As autumn puts an end to the growing season in preparation for winter in the north
temperate zones of the United States, I have been thinking about many things. I feel rather like my
mind is having a fantastic tea party such as Alice encountered in her trip down the rabbit hole. I
want very much to make you aware of the joys of orchid hunting in the wild. I also want to describe
these experiences in all their numerous sensory dimensions that I have experienced this season. I
hope I don‘t overload my word brush with too much information and overwork your receptors as I
flit from talk of ‗cabbages‘ to talk of ‗kings‘.
When I have a chance to go exploring for wild orchids I enter into the experience on many
levels of perception. I use my senses broadly, almost unaware that, until I have a chance to stop, sort
out, and reflect on my experiences, that I am accumulating so many sensations. Let me try to
describe an expedition that included a hike along a mountain trail.
The day is bright, sunny, and warm. Shortly after leaving home we stopped at our favorite
roadside site for Spiranthes ochroleuca, the yellow ladies‘-tresses. We are rewarded with several hundred
creamy yellow spikes along a wide stretch. It is a feast for my eyes to see the natural tapestry of the
plants and the elegant display of orchids tucked everywhere on the hillside. There are asters,
silverrods, ferns, and lycopods intertwining among the grasses and the Spiranthes, as they lift their
spiral spikes regally, to accent the scene. We are off to a flying start!
Spiranthes ochroleuca on a hillside in Carroll Co., NH
122
We have just traveled two and a half hours by car to reach the next spot. It is deep in the
northern NH mountains about five miles up a dusty dirt road that is heavily shaded with the forest
that seems to close in around us. As we near the end of the road the landscape opens up, thanks to
logging, and we feel the heat of the sun and squint in the brightness. This area looks promising for
our quest today, Spiranthes casei, Case‘s ladies‘-tresses. We know the flowering time is right and we
know that the soils are right and this spot has the open bankings that the orchids like to grow in so
we are encouraged to explore the area.
We traversed the area that has been logged, which opened up the shady roadsides to the full
sun. We did not find any orchids in that particular area. It was too dry and too open. That did not
dampen our enthusiasm for there were more banks down the road to explore.
Almost as soon as we reached these banks we were rewarded with great quantities of
Spiranthes cernua, nodding ladies‘-tresses, no S. casei among them. Working up the roadside we had
success. There were five plants coming into flower – a new record for this area.
At the bottom of the road a breathtaking vista from a wooden bridge beckoned me. There
had been several wooden bridges over brooks and small streams but this one gave me a view of two
tumbling streams, rushing together below me. There was a display of joe-pye weed and goldenrods
framing the nearer edges of the streams. The waters sang incessantly in my ears. A fritillary butterfly
lit on my hand for a brief inspection and lifted off in search of a more delicious quarry. A hiking trail
in the deep shade beside the stream banks looked intriguing – a welcome relief after the hot, sunny
explorations along the road.
We had other orchid habitatas to seek out and we had heard there were green-flowered rein
orchids along the trailside. Even though they would be in fruit it was an exciting search and could be
just as rewarding as finding flowering plants.
The trail was wet from yesterday‘s rains and slippery under foot. Large rocks, like stepping
stones, helped us navigate these areas but you felt like an awkward ballet dancer or inept mountain
goat as you tried to keep your balance. The trail side had several clumps of Cypripedium acaule, pink
lady‘s-slippers, that must have been spectacular earlier in the season. At the edge of a runoff that cut
diagonally across the trail a striking lavender-purple epipactis stood guard proudly. A little search
yielded several more plants and down the slope small purple fringed orchids were discovered which,
like the lady‘s-slippers, would have been glorious earlier.
Our climb took us further away from the streams until their rushing waters were diminished
to a soft murmur. We finally tired and retraced our steps back to our starting point – a long but
memorable trek, even though we didn‘t find the green rein orchids.
As the day stretched toward evening we continued our explorations for Spiranthes. We
checked several promising roadsides coming up empty-handed. A state park gave us a few Spiranthes
ochroleuca. It was an open hilltop with serpentine outcrops in rich deep gray green colors, the field
was dotted with tasty wild blueberries and grasses with the Spiranthes hiding there and there among
them.
As it grew darker it was harder to pick up indications of white spikes along our drive. Our
fleeting glimpses playing teasingly on our weary eyes promised us more adventure in discovery when
we return to the area another day.
Later in the darkness of my bedroom after such a day in the field my tired muscles relax
luxuriantly, but as I close my eyes, the vivid images of the hillsides spring to life. I see the patches of
moss, green haircap, gray green reindeer lichen, trailing fingers of lycopods with their stiff upright
fruiting stems intermingled with the delightfully jaunty, creamy white spikes of the Spiranthes. The
entire scene unfurls across my mind‘s eye in magnificent panoply. I see the bright colors of the
123
fritillary and can recall the gentle kiss of its touch on my hand. I can recall the sounds of the rushing
cataracts as they tumble playfully over the rocks, although I can no longer hear them in the silence
of my bedroom. I remember the cool shade of the forest trail and the warm caress of the sun that
heated me and made me blink in its brightness. My nose recalls the dust of the dirt road and the
damp earthy smell of the forest trail. The sweet taste of the wild blueberries lingers in my thoughts
and the lure of another day‘s adventuring lulls my tired senses into a deep, refreshing sleep.
I hope most of your expeditions expand your horizons, excite all of your senses and fulfill
your quests. It‘s called being alive to all of the possibilities.
Your Slow Empiricist
124
RECENT ARTICLES
RECENT ARTICLES OF INTEREST IN OTHER JOURNALS

Orchids magazine form the American Orchid Society often has one or more articles each month
about North American natives. Check out their website for issue contents. http://magazine-
directory.com/Orchids.htm
SOUTHEASTERN NATURALIST 7(4):571-580. 2008

Host-tree Selection by an Epiphytic Orchid, Epidendrum magnoliae Muhl. (Green Fly
Orchid), in an Inland Hardwood Hammock in Georgia
Bradley J. Bergstrom and Richard Carter
ABSTRACT: We characterized the tree community of a mesic hardwood hammock in south-central
Georgia as an oak-pine-hickory forest, with Liquidambar styraciflua (Sweetgum), Magnolia grandiflora
(Southern Magnolia), and Ilex opaca Ait. (American Holly) as subdominants. We surveyed this forest
for colonies of the most northerly distributed epiphytic orchid in the Western Hemisphere,
Epidendrum magnoliae (Green Fly Orchid), and recorded the species and trunk diameter of 112 host
trees (phorophytes) as well as the height and size of each orchid colony. We calculated a selectivity
index (SI) to compare phorophyte frequency with availability, based on a point-transect survey.
Green Fly Orchid occurred on 8 species of hardwood trees, but had a strong preference for
Southern Magnolia as a host and a moderately strong preference for Quercus virginiana (Live Oak).
Host trees were much larger (presumably older) than the average of available trees, and that effect
was strongest for the most preferred host. Orchid colonies also occupied significantly greater areas
on individual Southern Magnolia than on other phorophytes. It is likely that old-growth Southern
Magnolia and Live Oak trees are critical to the viability of this population of Green Fly Orchid,
which is rare in inland forests in Georgia. In addition to being the most persistent epiphyte
substrates in this environment, their broadleaf evergreen canopies—which would be especially true
of Southern Magnolia —may provide the most favorable microclimates in terms of shade, humidity,
and frost protection.
AMERICAN JOURNAL OF BOTANY. 95:498-505. 2008

Relationships and evolution of matK in a group of leafless orchids (Corallorhiza and
Corallorhizinae; Orchidaceae: Epidendroideae)
John V. Freudenstein and Diana M. Senyo
ABSTRACT
Corallorhizinae are a small group of Old and New World temperate orchids of which a core
monophyletic group comprises Govenia, Cremastra, Aplectrum, Oreorchis and the leafless Corallorhiza, and
which according to phylogenetic analysis of nuclear ITS and plastid matK sequences, are related in
this way: (Govenia (Cremastra (Aplectrum (Oreorchis (Corallorhiza))))). This hypothesis is consistent with
the progressive deletion of the trnK intron and matK ORF. Frameshift-resulting indels yield a
predicted loss of translation for the critical "domain X" region of matK and are evidence that matK is
a probable pseudogene in Aplectrum, Oreorchis, and Corallorhiza. Within Corallorhiza, a previous
hypothesis based on plastid DNA restriction site analysis is confirmed, with the thickened-labellum
C. striata group being sister to the thin-labellum remainder of the genus, within which the
circumboreal C. trifida is sister to the remainder, which then comprise two further sister groups: C.
maculata + C. bulbosa + C. mertensiana and C. odontorhiza + C. wisteriana. A close relationship between
C. striata and the recently described Appalachian C. bentleyi is shown; in particular, C. bentleyi is more
closely allied to a southern Mexican population of C. striata than it is to northern North American C.
striata populations, suggesting that two lineages, each with Mexican and northern North American
populations, exist within the C. striata group.
125
RECENT ARTICLES
ANNALS OF BOTANY 102: 783-793. 2008.

Asymbiotic Germination Response to Photoperiod and Nutritional Media in Six
Populations of Calopogon tuberosus var. tuberosus (Orchidaceae): Evidence for Ecotypic
Differentiation
Philip J. Kauth, Michael E. Kane, Wagner A. Vendrame and Carrie Reinhardt-Adams
Background and Aims: Ecotypic differentiation has been explored in numerous plant species, but
has been largely ignored in the Orchidaceae. Applying a specific germination protocol for
widespread seed sources may be unreliable due to inherent physiological or genetic differences in
localized populations. It is crucial to determine whether ecotypic differentiation exists for restoration
and conservation programmes. Calopogon tuberosus var. tuberosus, a widespread terrestrial orchid of
eastern North America, is a model species to explore ecotypic differences in germination
requirements, as this species occupies diverse habitats spanning a wide geographical range.
Methods: Mature seeds were collected from south Florida, north central Florida, three locations in
South Carolina, and the upper Michigan peninsula. Effects of three photoperiods (8/16, 12/12, 16/8
h L/D) were examined on asymbiotic in vitro seed germination and seedling development of C.
tuberosus. Germination and early development was monitored for 8 weeks, while advanced
development was monitored for an additional 8 weeks. In an additional experiment, asymbiotic seed
germination and development was monitored for 8 weeks on six culture media (BM-1 terrestrial
orchid medium, Knudson C, Malmgrem, half-strength MS, P723, and Vacin and Went). A
tetrazolium test for embryo viability was performed.
Key Results: Short days promoted the highest germination among Florida populations, but few
differences among photoperiods in other seed sources existed. Different media had little effect on
the germination of Michigan and Florida populations, but germination of South Carolina seeds was
higher on media with higher calcium and magnesium. Tetrazolium testing confirmed that South
Carolina seeds exhibited low viability while viability was higher in Florida seeds. Seed germination
and corm formation was rapid in Michigan seeds across all treatments. Michigan seedlings allocated
more biomass to corms compared with other seed sources.
Conclusions: Rapid germination and corm formation may be a survival mechanism in response to a
compressed growing season in northern populations. Ecotypic differentiation may be occurring
based on seed germination and corm formation data.
SYSTEMATIC BOTANY 34(3): 496–504. 2009.

Patterns of Morphological and Plastid DNA Variation in the Corallorhiza striata Species
Complex (Orchidaceae)
Craig F. Barrett and John V. Freudenstein
ABSTRACT: Corallorhiza striata is a wide-ranging, morphologically variable, mycoheterotrophic
species complex distributed across North America. Objectives of this study were to assess
relationships and test validity of previously delimited varieties of C. striata, including the recently
described C. bentleyi . Two plastid DNA regions were sequenced for individuals from several
populations across North America, identifying four major clades. The large-flowered C. striata var.
striata (northern U.S.A., southern Canada) was sister to the smaller-flowered var. vreelandii
(southwestern U.S.A., Mexico), and these were sister to a Californian clade with relatively
intermediate-sized flowers. C. striata var. involuta (Mexico) and the endangered C. bentleyi (eastern
U.S.A.) shared a close relationship, sister to the remaining C. striata . Principal Components Analysis
and Nonparametric Multivariate Analysis of Variance on nine quantitative morphological characters,
using plastid DNA clades as independent variables, demonstrated strong correlations between
126
RECENT ARTICLES
molecular and morphological groupings. Morphological analyses supported differentiation of both

C. striata var. involuta and C. bentleyi relative to all other accessions of C. striata , suggesting their
recognition as separate species; these findings will have future implications for conservation. The
biogeographic scenario was more complex than previously thought, with members of two major
plastid DNA lineages ( C. bentleyi /var. involuta and the remaining C. striata ) existing in Mexico and
U.S.A./Canada. These findings contribute to a burgeoning body of data on poorly studied North
American plant distributions extending into southern Mexico.
LANKESTERIANA 8: 105-112. 2009.

Vegetative anatomy of Calypsoeae (Orchidaceae)
William Louis Stern and Barbara S. Carlsward
ABSTRACT: Calypsoeae represent a small tribe of anatomically little-known orchids with a wide
distribution in the Western Hemisphere. Leaves are present in all genera, except Corallorhiza and
Wullschlaegelia both of which are subterranean taxa. Stomata are abaxial (ad- and abaxial in Aplectrum)
and tetracytic (anomocytic in Calypso). Fiber bundles are absent in leaves of all taxa examined except
Govenia tingens. Stegmata are present in leaves of only Cremastra and Govenia. Roots are velamentous,
except in filiform roots of Wullschlaegelia. Vegetative anatomy supports a relationship between
Wullschlaegelia and Corallorhiza but does not support the grouping of winter-leaved Aplectrum and
Tipularia nor proposed groupings of genera based on pollinarium features.
PLANT CELL, TISSUE AND ORGAN CULTURE 96: 235-243. 2009.

Asymbiotic seed germination and in vitro seedling development of Cyrtopodium
punctatum: a propagation protocol for an endangered Florida native orchid.
Daniela Dutra, Michael E. Kane, and Larry Richardson
ABSTRACT: Cyrtopodium punctatum Lindley is an endangered epiphytic orchid restricted in the United
States to southern Florida. Due to its ornamental value, the species was extensively collected from
the wild during the past 100 years. Today, only a few plants remain in protected areas. As part of a
conservation plan for the species, procedures for asymbiotic seed germination were developed. Five
asymbiotic orchid seed germination media (PhytoTechnology Orchid Seed Sowing Medium,
Knudson C, Malmgren Modified Terrestrial Orchid Medium, Vacin & Went Modified Orchid
Medium, and _-strengh Murashige & Skoog) were examined for their effectiveness in promoting
seed germination and protocorm development under a 16/8 h L/D photoperiod and dark (0/24 h
L/D). The influence of photoperiod on growth and development was also examined. Seeds were
germinated under a 16/8 h, 12/12 h, 8/16 h L/D photoperiod, at 25 ± 3_C and allowed to develop
in vitro for 10 weeks. After 10 weeks, developing seedlings were transferred to Sigma Phytatrays and
returned to their assigned photoperiod treatments for continued seedling development for an
additional 15 weeks. Highest germination occurred in 0/24 h L/D on PhytoTechnology Orchid
Seed Sowing Medium and seedlings displayed more advanced development when cultured under
16/8 h L/D photoperiod after 15 weeks in Phytatrays. Thirty-five week old seedlings potted in
coconut husk growing medium exhibited 90% survival following 5 weeks acclimatization to
greenhouse conditions. This asymbiotic seed germination protocol for C. punctatum will facilitate
future reintroduction projects involving this endangered species.
NATIVE ORCHID CONFERENCE JOURNAL 6(1): 17-20. 2009

A Yellow Form of Dichromanthus michuacanus (Llave & Lex.) Salazar & Soto Arenas in
Arizona Ronald A. Coleman
Narrative and description of Dichromanthus michuacanus forma armeniacus.
127
Kieffer: PLATANTHERA ×CANBYI
REDISCOVERIES OF PLATANTHERA ×CANBYI ON THE

EASTERN SHORE OF MARYLAND
Chelsea Kieffer
EDITOR’S NOTE: Just before going to press I had an email for Chelsea regarding the recent discovery of Platanthera
xcanbyi on the eastern shore of Maryland. She noted that she had found the same hybrid along with the parents in 2004
in what is perhaps the same locale. This is her account and terrific photos of the three taxa.
A Washington Post article recently caused quite a commotion for orchid lovers when it
reported that a rare hybrid of White Fringed (Platanthera blephariglottis) and Crested Yellow Fringed
(Platanthera cristata) Orchids was rediscovered on the Eastern Shore of Maryland as a result of a
controlled burning. The orchid, known as Platanthera ×canbyi, was last seen 18 years ago according to
the article.
The managers of Maryland's largest private nature preserve burned 240 acres this spring in the
hopes of triggering the growth of long-dormant native plants. The plan worked, producing a rare
hybrid orchid that has only been found in Maryland once before -- 18 years ago. -THE
WASHINGTON POST 08/12/09
My parents and I scoped out the Nassawango area of Maryland‘s Eastern Shore back in 2004
at the suggestion of a friend who had knowledge of the flora of that area. On July 3, 2004 I found
numerous plants with flower buds showing some color. These included tall (28‖ +/-), robust plants
with pale yellow flower buds and short (4-10‖), not as vigorous plants with yellow orange flower
buds. I knew I would have to revisit this site in a couple of weeks, if I wanted to see them in full
bloom.
On our return visit on July 17, 2004, I photographed several individuals of Platanthera
blephariglottis and Platanthera cristata, as well as both species growing side by side in one location.
Then, within a few yards I found and photographed a single plant exhibiting size and flower color
characteristics combining those of the two species, and suspected that this was a natural hybrid of
the two species. I consulted with Chapman‘s Orchids of the Northeast and concluded that this plant was
likely Platanthera ×canbyi!
I had no idea that this hybrid had not been found for 18 years. As of this writing I have
initiated contact with the land manager of the Nassawango Creek Preserve in order to compare
notes.
128
Kieffer: PLATANTHERA ×CANBYI
Platanthera blephariglottis (left) and P. cristata

(right)
Platanthera ×canbyi
Photos by Chelsea Kieffer
129
BOOK REVIEWS
BOOK REVIEWS
Ireland’s Wild Orchids

by Brendan Sayles and Susan Sex
Rarely does a book come along that is virtually perfect. Field Guide to the Orchids of Ireland is
this book After receiving copies of both the spiral bound and leather bound editions there is next to
nothing I can say other than ‗what a perfect field guide. The text is complete and easy to understand,
the keys comprehensive, and the photos and watercolors illustrate all of the species beautifully. The
spiral binding is durable and the pages are on heavy-weight paper. For the collector the limited
edition leather bound version with a slip case is a must for any library. The price of the spiral bound
edition is more than reasonable and, although a bit pricy, the leather edition is worth every penny
and contain additional water colors sketches. PMB
From the authors‘ website:

―The field guide is in the manner of the notebooks and sketchpads used by artists and writers for
more than a century. Designed to fit the pocket, be shower proof and open flat in the field, the
standard copy has an inside pocket, space for field notes and photographs and a strong elastic for
holding the book closed when required. This book contains 119 pages, filled with paintings by
Susan, text notes and new accessible distinguishing keys to the species from Brendan and
photographs from contributing orchideers such as Leueen Hill from the Burren to Jackie O‘Connell
from the Midlands. It promises to make the mysteries of orchid identification easier whilst being a
beautiful book in its own right.‖
For more information and purchase details see www.orchidireland.ie
130
BOOK REVIEWS
Wild Orchids in the Burren

Pat O‘Reilly and Sue Parker
First Nature Guides
Full color, 67 pages, paperbound 6 x 8.25‖
ISBN 0-9549554-8-X 2007
For prices and shipping contact enquires@first-nature.com
This excellent little paperback present striking color photos of both the orchids and the fascinating
region known as ‗The Burren‘ in southwestern Ireland. Far from being the ‗barrens‘ of the name,
this area is rich in rare and common plants and especially in orchids. Each species found there is
treated with several color photos and information on flowering period, habitat, geographic area, and
degree of rarity as well as hybrids and color forms. Also included is a map of existing trails for those
who visit and wish to hike the region. Highly recommended. PMB
131
NEW TAXA
NEW COMBINATIONS IN VOLUME 15(2)

Blephariglottis albiflora Raf. forma holopetala (Lindl. ) Baumbach & P.M. Brown comb. nov.
Blephariglottis cristata Raf. forma straminea (P.M. Brown) Baumbach & P.M. Brown comb. nov.
Blephariglottis grandiflora (Lindley) Rydb. forma albiflora (Rand & Redf. ) Baumbach & P.M.
Brown comb. nov.
Blephariglottis grandiflora (Bigelow) Rydb. forma bicolor (P.M. Brown) Baumbach & P.M. Brown
comb. nov.
Blephariglottis grandiflora (Lindl.) Rydb. forma carnea (P.M. Brown) Baumbach & P.M. Brown
comb. nov.
Blephariglottis grandiflora (Bigelow) Rydb. forma mentotonsa (Fernald ) Baumbach & P.M. Brown
comb. nov.
Blephariglottis peramoena (A. Gray) Rydb. forma doddsiae (P.M. Brown) Baumbach & P.M. Brown
comb. nov.
Blephariglottis psycodes (L.) Rydb. forma albiflora (R. Hoff.) Baumbach & P.M. Brown comb. nov.
Blephariglottis psycodes (L.) Rydb. forma fernaldii (J. Rousseau & Rouleau) Baumbach & P.M.
Brown comb. nov.
Blephariglottis psycodes (L.) Rydb. forma ecalcarata (Bryan) Baumbach & P.M. Brown comb. nov.
Blephariglottis psycodes (L.) Rydb. forma rosea (P.M. Brown) Baumbach & P.M. Brown comb. nov.
Blephariglottis psycodes (L.) Rydb. forma varians (Bryan) Baumbach & P.M. Brown comb. nov.
Limnorchis aquilonis (Sheviak) Rebrist. & Elven forma alba (Light) P.M. Brown, S.L. Stewart & R.
Gamarra comb. nov.
Limnorchis brevifolia (Greene) Rydb. Forma alba (P.M. Brown) P.M. Brown, S.L. Stewart & R.
Gamarra comb. nov.
Limnorchis dilatata (Pursh) Rydb. var. leucostachys (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra
comb. nov.
Limnorchis holochila (Hillebr.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis limosa (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis tescamnis (Sheviak & W.F. Jenn.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis tipuloides (L. f.) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis tipuloides ( L. f.) P.M. Brown, S.L. Stewart & R. Gamarra var. behringiana (Rydb.) P.M.
Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis yosemitensis (Colwell, Sheviak & P.E. Moore) P.M. Brown, S.L. Stewart & R. Gamarra
comb. nov.
Limnorchis ×correllii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×estesii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×evansiana (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×folsomii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×kelleyi (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×lassennii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
Limnorchis ×smithii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra comb. nov.
132
133
134
135
FROM THE SWAMPS OF SOUTH FLORIDA TO THE WILDS OF NORTHERN ALASKA….
TO WINDSWEPT NEWFOUNDLAND AND THE BIG BEND OF WEST TEXAS
WILD ORCHIDS….
from the University Press of Florida
by Paul Martin Brown & Stan Folsom
Ordering information from University Press of Florida www.upf.com or 1-800-226-3822

or for signed and inscribed copies from the authors at naorchid@aol.com
136
or directly from the author at naorchid@aol.com
137

June 2009 North American Native Orchid Journal

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June 2009 North American Native Orchid Journal

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

Paul Martin Brown, Editor

Scott L. Stewart, PhD. Associate Editor

RECENT PUBLICATIONS REGARDING PLATANTHERA AND ALLIED

Paul Martin Brown, Scott L. Stewart and authors

MOLECULAR PHYLOGENETICS AND MORPHOLOGICAL REAPPRAISAL OF THE

Delimitation and relationships of genera and sections in the Platanthera clade

The Galearis clade: Galearis s.l.

The Platanthera clade: section Limnorchis

Because of the exceptional taxonomic instability surrounding the delimitation of P. hyperborea

Sections Lacera and Tulotis

Sections Lysiella and Lysias

(a) Former genera Amerorchis, [Aceratorchis and Chondradenia]

Die Gattungen Platanthera L.C. RICH. und Blephariglottis RAF.

New combinations for formae are proposed here:

The genus Blephariglottis in North America as treated by Baumbach and Lückel.

Blephariglottis psycodes (L.) Rydb. Blephariglottis ×canbyi (Ames) W. Stone

SEED MICROMORPHOLOGY SUPPORTS THE SPLITTING OF

The Genus Limnorchis in the United States and Canada.

What does all of this mean?

LITERATURE CITED IN EXTRACTIONS FROM BATEMAN ET AL.

LITERATURE CITED IN EXTRACTIONS FROM GAMARRA ET AL.

LITERATURE CITED IN EXTRACTIONS FROM BAUMBACH AND LÜCKEL

Luer, C. A. (1972): The native Orchids of Florida: 136-151

Blephariglottis chapmanii Blephariglottis lacera Blephariglottis pallida

Blephariglottis praeclara Blephariglottis peramoena Blephariglottis ×channellii

Blephariglottis ×lueri Blephariglottis ×andrewsii

Limnorchis brevifolia Limnorchis limosa

Limnorchis dilatata Limnorchis yosemitensis Limnorchis holochila L. hyperborea (Greenland)

THE AMERICAN SPECIES OF LIMNORCHIS

The following is a facsimile reprint of Rydberg‘s original work on

WILD ORCHIDS IN THE 21ST CENTURY:

Wild Orchids of the Northeast: New England,

Field Guide to the Wild Orchids of Texas

Each of the 51 Quebec orchids

Limited printing of 2000 copies

2007 2007 2008

The first 5 laminated

Genera Orchidacearum, Volume 5:

Edited by Alec M Pridgeon, Phillip J Cribb, Mark W Chase

…..OF CABBAGES AND KINGS

Spiranthes ochroleuca on a hillside in Carroll Co., NH

RECENT ARTICLES OF INTEREST IN OTHER JOURNALS

SOUTHEASTERN NATURALIST 7(4):571-580. 2008

AMERICAN JOURNAL OF BOTANY. 95:498-505. 2008

ANNALS OF BOTANY 102: 783-793. 2008.

SYSTEMATIC BOTANY 34(3): 496–504. 2009.

molecular and morphological groupings. Morphological analyses supported differentiation of both

LANKESTERIANA 8: 105-112. 2009.

PLANT CELL, TISSUE AND ORGAN CULTURE 96: 235-243. 2009.

NATIVE ORCHID CONFERENCE JOURNAL 6(1): 17-20. 2009

REDISCOVERIES OF PLATANTHERA ×CANBYI ON THE

Platanthera blephariglottis (left) and P. cristata

Photos by Chelsea Kieffer

Ireland’s Wild Orchids

From the authors‘ website:

Wild Orchids in the Burren

NEW COMBINATIONS IN VOLUME 15(2)

Ordering information from University Press of Florida www.upf.com or 1-800-226-3822

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