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
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 THE AMERICAN SPECIES OF LIMNORCHIS 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
Volume 15 (2) 2009

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

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 

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
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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 sistergroup 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
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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
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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
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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.
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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.

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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.
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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

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Blephariglottis peramoena (A. Gray) Rydb. forma doddsiae (P.M. Brown) Baumbach & P.M. Brown comb. nov.
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 & P.M. Brown comb. nov.
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. forma carnea (P.M. Brown) Baumbach & P.M. Blephariglottis albiflora Raf.
Syns: Platanthera blephariglottis (Willd.) Lindl., Habenaria blephariglottis (Willd.) Hooker

forma holopetala (Lindl.) Baumbach & P.M. Brown

Blephariglottis chapmanii Small

Syns: Platanthera chapmanii (Small) Luer, Habenaria xchapmanii (Small) Ames

Brown forma mentotonsa (Fernald ) Baumbach & P.M. Brown Blephariglottis longicornis Raf. Syns.: Platanthera

integrilabia (Correll) Luer, Habenaria blephariglottis var. integrilabia Correll, Habenaria correlliana Cronquist

Blephariglottis ciliaris (L.) Rydb.
Syns.: Platanthera ciliaris Lindl., Habenaria ciliaris (L.) R. Brown

Blephariglottis lacera (Michx.) Farw.
Syns.: Platanthera lacera (Michx.) G. Don, Habenaria lacera Michx. R. Brown

Blephariglottis conspicua (Nash) Small
Syns.: Platanthera conspicua (Lindl.) P.M. Brown, Habenaria conspicua Nash (not treated in Baumbach & Lückel)

Blephariglottis leucophaea (Nutt.) Farw.
Syns.: Platanthera leucophaea (Nutt.) Lindl., Habenaria leucophaea (Nutt.) Gray

Blephariglottis cristata Raf.

Syns.: Platanthera cristata Michx., Habenaria cristata (Michx.) R. Brown

Blephariglottis pallida (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera pallida P.M. Brown

forma straminea (P.M. Brown) Baumbach & P.M. Brown

Blephariglottis peramoena (A. Gray) Rydb.

Blephariglottis grandiflora (Bigelow) Rydb.

Syns.: Platanthera peramoena (A. Gray) A. Gray, Habenaria peramoena A. Gray

Syns.: Platanthera grandiflora (Bigelow) Lind., Habenaria grandiflora (Bigelow) Torrey ex L.C. Beck forma albiflora (Rand & Redf. ) Baumbach & P.M.

forma doddsiae (P.M. Brown) Baumbach & P.M. Brown

Brown forma bicolor (P.M. Brown) Baumbach & P.M. Brown

Blephariglottis praeclara (Sheviak & Bowles) Baumb. & Lueckel

Syns.: Platanthera praeclara Sheviak & Bowles, Habenaria leucophaea A. Gray var. praeclara (Sheviak & Bowles) Cronquist

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Blephariglottis psycodes (L.) Rydb.
Syns.: Platanthera psycodes (L.) Lindl., Habenaria psycodes (L.) Sprengl.

Blephariglottis ×canbyi (Ames) W. Stone
Syns.: Platanthera ×canbyi (Ames) Luer, Habenaria ×canbyi Ames

forma albiflora (R. Hoff.) Baumbach & P.M. Brown forma fernaldii (J. Rousseau & Rouleau) Baumbach & P.M. Brown forma ecalcarata (Bryan) Baumbach & P.M. Brown forma rosea (P.M. Brown) Baumbach & P.M. Brown forma varians (Bryan) Baumbach & P.M. Brown

Blephariglottis ×channellii (Folsom) Baumb. & Lueckel
Syn.: Platanthera ×channellii Folsom

Blephariglottis ×enigma (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera ×enigma P.M. Brown

Blephariglottis shriveri (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera shriveri P.M. Brown

Blephariglottis ×hollandiae (Catling & Brownell) Baumb. & Lueckel
Syn.: Platanthera ×hollandiae Catling & Brownell

Blephariglottis ×andrewsii (M. White) House,
Blephariglottis ×apalachicola (P.M. Brown & S. Stewart) Baumb. & Lueckel

Blephariglottis ×keenanii (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera ×keenanii P.M. Brown

Syns.: Platanthera ×andrewsii (M. White) Luer, Habenaria ×andrewsii M. White Syn.: Platanthera ×apalachicola P.M. Brown & S. Stewart

Blephariglottis ×lueri (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera ×lueri P.M. Brown

Blephariglottis ×beckneri (P.M. Brown) Baumb. & Lueckel
Syn.: Platanthera ×beckneri P.M. Brown

Blephariglottis ×osceola (P.M. Brown & S. Stewart) Baumb. & Lueckel
Syn.: Platanthera ×osceola P.M. Brown & S. Stewart

Blephariglottis ×bicolor Raf.

Blephariglottis ×reznicekii (Catling, Brownell & Allen) Baumb. & Lueckel
Syn.: Platanthera ×reznicekii Catling, Brownell & Allen

Syn.: Platanthera ×bicolor (Raf.)Luer

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 socalled ‗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
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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
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‗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

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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. Gamarra
Syns.: Platanthera dilatata (Pursh) Lindl. var. albiflora (Cham.) Ledeb., Habenaria dilatata var. albiflora (Cham.) Correll

Limnorchis brevifolia (Greene) Rydb.

Syns.: Platanthera brevifolia (Greene) Kranzl., Habenaria brevifolia Greene

Limnorchis dilatata (Pursh) Rydb. var. leucostachys (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra
Syns.: Limnorchis leucostachys (Lindl.) Rydb., Platanthera dilatata (Pursh) Lindl. var. leucostachys (Lindl.) Luer, Habenaria dilatata var. leucostachys (Lindl.) Ames

forma alba (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra

Limnorchis chorisiana (Cham.) J.P. Anderson

Syns. Platanthera chorisiana (Cham.) Reich. f., Habenaria chorisiana Cham.

Limnorchis huronensis (Nutt.) Rydbg.
Syns.: Platanthera huronensis (Nutt.) Lindl., Habenaria huronensis (Nutt.) Spreng.

Limnorchis convallariifolia (Fischer ex Lindl.) Rydb.
Syns.: Platanthera convallariifolia Fischer ex Lindl., Habenaria convallariifolia (Lindl.) B. Boivin

Limnorchis hyperborea (L.) Rydb.

Limnorchis dilatata (Pursh) Rydb. var. dilatata
Syns.: Platanthera dilatata (Pursh) Lindl. var. dilatata, Habenaria dilatata (Pursh) Hooker

Syns.: Platanthera hyperborea (L.) Lindl., Habenaria hyperborea (L.) R. Br. ex W.T. Aiton (known only from Greenland and Iceland)

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Limnorchis holochila (Hillebr.) P.M. Brown, S.L. Stewart & R. Gamarra
Syns.: Habenaria holochila Hillebr., Platanthera holochila (Hillebr.) Kraenzl. (known only from Hawaii)

Limnorchis viridiflora ( Cham. ) Rydb.
Syns: Platanthera hyperborea var. viridiflora (Cham.) Luer, Habenaria borealis Cham. var. viridiflora Cham. (known from western Alaska per Bateman et al.)

Limnorchis limosa (Lindl.) P.M. Brown, S.L. Stewart & R. Gamarra
Syns.: Platanthera limosa Lindl., Habenaria limosa (Lindl.) Hemsl.

Limnorchis yosemitensis (Colwell, Sheviak & P.E. Moore) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera yosemitensis Colwell, Sheviak & P.E. Moore

Limnorchis purpurascens Rydb.

Limnorchis zothecina (L.C. Higgins & S.L. Welsh) W.A. Weber
Syns.: Platanthera zothecina (L.C. Higgins & S.L. Welsh) Kartesz & Gandhi, Habenaria zothecina L.C. Higgins & S.L. Welsh

Syns.: Platanthera purpurascens (Rydb.) Sheviak & W.F. Jenn., Habenaria purpurascens (Rydb.) Tidestr. in Tidestr. & Kittell

Limnorchis sparsiflora (S. Watson) Rydb.

Syns.: Platanthera sparsiflora (S. Watson) Schltr., Habenaria sparsiflora S. Watson

Limnorchis ×correllii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×correllii W.J. Schrenck

Limnorchis stricta (Lindl.) Rydb.

Syns.: Platanthera stricta Lindl., Habenaria stricta (Lindl.) Rydb.

Limnorchis ×estesii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×estesii W.J. Schrenck

Limnorchis tescamnis (Sheviak & W.F. Jenn.) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera tescamnis Sheviak & W.F. Jenn.

Limnorchis ×evansiana (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×evansiana P.M. Brown

Limnorchis ×folsomii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×folsomii P.M. Brown

Limnorchis tipuloides (L. f.) P.M. Brown, S.L. Stewart & R. Gamarra
Syns: Platanthera tipuloides (L. f. ) Lindl., Habenaria tipuloides (L. f. ) Bentham

Limnorchis ×kelleyi (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×kelleyi P.M. Brown

Limnorchis tipuloides ( L. f.) P.M. Brown, S.L. Stewart & R. Gamarra var. behringiana (Rydb.) P.M. Brown, S.L. Stewart & R. Gamarra
Syns.: Limnorchis behringiana Rydb., Platanthera tipuloides (L. f.) Lindl. var. behringiana (Rydb.) Hultén

Limnorchis ×lassennii (W.J. Schrenck) P.M. Brown, S.L. Stewart & R. Gamarra
Syn.: Platanthera ×lassenii W.J. Schrenk

Limnorchis ×smithii (P.M. Brown) P.M. Brown, S.L. Stewart & R. Gamarra
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.
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 

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 LITERATURE CITED IN EXTRACTIONS FROM BATEMAN ET AL. Ackerman JD. 1977. Biosystematics of the genus Piperia Rydb. (Orchidaceae). Botanical Journal of the Linnaean Society 75: 245–270. Ackermann JD, Morgan R. 2002. Piperia Rydberg. In: Flora of North America Editorial Committee. eds. Flora of North America North of Mexico 26. Oxford: Oxford University Press, 571–577. Bateman RM. 2001. Evolution and classification of European orchids: insights from molecular and morphological characters. Journal Europaischer Orchideen 33: 33–119. Bateman RM. 2005. Circumscribing and interpreting closely related orchid species: Platanthera, Dactylorhiza and the crucial role of mutation. Journal of the Hardy Orchid Society 2: 104–111. Bateman RM, Denholm I. 1983. A reappraisal of the British and Irish dactylorchids, 1. The tetraploid marsh-orchids. Watsonia 14: 347–376. Bateman RM, Rudall PJ. 2006. The good, the bad, and the ugly: using naturally occurring terata to distinguish the possible from the impossible in orchid floral evolution. Aliso 22: 481–496. Bateman RM, Sexton R. 2008. Is spur length of Platanthera species in the British Isles adaptively optimized or an evolutionary red herring? Watsonia 28: 1–21. Bateman RM, Pridgeon AM, Chase MW. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 2. Infrageneric relationships and taxonomic revision to achieve monophyly of Orchis sensu stricto. Lindleyana 12: 113–141. Bateman RM, Hollingsworth PM, Preston J, Luo Y-B, Pridgeon AM, Chase MW. 2003. Molecular phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidaceae). Botanical Journal of the Linnaean Society 142: 1–40. Bateman RM, Rudall PJ, James KE. 2006. Phylogenetic context, generic affinities and evolutionary origin of the enigmatic Balkan orchid Gymnadenia frivaldii Hampe ex Griseb. Taxon 55: 107–118. Box MS, Bateman RM, Glover BJ, Rudall PJ. 2008. Floral ontogenetic evidence of repeated speciation via paedomorphosis in subtribe Orchidinae (Orchidaceae). Botanical Journal of the Linnaean Society 157: 429– 454. Catling PM, Catling VR. 1991. A synopsis of breeding systems and pollination in North American orchids. Lindleyana 6: 187–210. Dressler RL. 1993. Phylogeny and classification of the orchid family. Portland: Timber Press. Gamarra R, Golan P, Herrera I, Ortunez E. 2008. Seed micromorphology supports the splitting of Limnorchis from Platanthera. Nordic Journal of Botany 26: 61–65. Hapeman JR, Inoue K. 1997. Plant–pollinator interactions and floral radiation in Platanthera (Orchidaceae). In: Givnish TJ, Sytsma KJ. eds. Molecular evolution and adaptive radiation. Cambridge: Cambridge University Press, 433– 454. Hultén E. 1968. Amerorchis nov. gen. Arkiv for Botanik Series 2, 7: 34–35. Hunt PF. 1971. Notes on Asiatic orchids: VI. Kew Bulletin 26: 171–185.

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Inoue KI. 1983. Systematics of the genus Platanthera (Orchidaceae) in Japan and adjacent regions with special reference to pollination. Journal of the Faculty of Science, University of Tokyo, Section III (Botany) 13: 285–374. Lauri R. 2007. The systematic and phylogenetic study of the subgenus Piperia (Orchidaceae) and closely related Platanthera. Botanical Society of America Abstracts, 241. Luer CA. 1975. The native orchids of the United States and Canada, excluding Florida. New York: New York Botanical Garden. Pridgeon AM, Bateman RM, Cox AV, Hapeman JR, Chase MC. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 1. Intergeneric relationships and polyphyly of Orchis sensu lato. Lindleyana 12: 89–109. Pridgeon AM, Cribb PJ, Rasmussen FN, Chase MC. eds . 2001. Genera Orchidacearum 2: Orchidoideae 1. Oxford: Oxford University Press. Rafinesque CS. 1833. Herbarium Rafinesquianum. Philadelphia. [3 parts, designated as extras of Atlantic Journal.] Reinhammar L-G. 1995. Evidence for two distinctive species of Pseudorchis (Orchidaceae) in Scandinavia. Nordic Journal of Botany 15: 469–481. Reinhammar L-G, Hedrén M. 1998. Allozyme differentiation between lowland and alpine populations of Pseudorchis albida s.lat. (Orchidaceae) in Sweden. Nordic Journal of Botany 18: 7–14. Rydberg PA. 1901a. Studies on the Rocky Mountain flora. Part 5. Bulletin of the Torrey Botanical Club 28: 266–284. Rydberg PA. 1901b. The North American species of Limnorchis and Piperia, north of Mexico. Bulletin of the Torrey Botanical Club 28: 605–643. Sheviak CJ. 1999. The identities of Platanthera hyperborea and P. huronensis, with the description of a new species from North America. Lindleyana 14: 193–203. Sheviak CJ. 2002. Platanthera Richard. In: Flora of North America Editorial Committee. eds. Flora of North America North of Mexico 26. Oxford: Oxford University Press, 551–571. Sheviak CJ, Bracht M. 1998. New chromosome number determinations in Platanthera. Native North American Orchid Journal 4: 168–172. Sheviak CJ, Catling PM. 2002. Galearis Rafinesque, Amerorchis Hultén. In: Flora of North America Editorial Committee. eds. Flora of North America North of Mexico 26. Oxford: Oxford University Press, 550–551. Wallace LE. 2002. Biological investigations in the genus Platanthera (Orchidaceae): conservation issues in Platanthera leucophaea and evolutionary diversification in section Limnorchis. Doctoral dissertation, Ohio State University. Wallace LE. 2003. Molecular evidence for allopolyploid speciation and recurrent origins in Platanthera huronensis (Orchidaceae). International Journal of Plant Sciences 164: 907–916. Wallace LE. 2004. A comparison of genetic variation and structure in the allopolyploid Platanthera huronensis and its diploid progenitors, Platanthera aquilonis and Platanthera dilatata (Orchidaceae). Canadian Journal of Botany 82: 244–252. Wallace LE. 2006. Spatial genetic structure and frequency of interspecific hybridization in Platanthera aquilonis and P. dilatata (Orchidaceae) occurring in sympatry. American Journal of Botany 93: 1001–1009. Wood JJ, Neiland RN. 2001a. Amerorchis. In: Pridgeon AM, Cribb PL, Rasmussen FN, Chase MW. eds. Genera Orchidacearum 2: Orchidoideae 1. Oxford: Oxford University Press, 245–247. Wood JJ, Neiland RN. 2001b. Galearis. In: Pridgeon AM, Cribb PL, Rasmussen FN, Chase MW. eds. Genera Orchidacearum 2: Orchidoideae 1. Oxford: Oxford University Press, 290–292. Wood JJ, Neiland RN. 2001c. Platanthera. In: Pridgeon AM, Cribb PL, Rasmussen FN, Chase MW. eds. Genera Orchidacearum 2: Orchidoideae 1. Oxford: Oxford University Press, 345–350. LITERATURE CITED IN EXTRACTIONS FROM GAMARRA ET AL. Ackermann, J. D. 1977. Biosystematics of the genus Piperia Rydb. (Orchidaceae). Bot. J. Linn. Soc. 75: 245270. Ames, O. 1910. Orchidaceae. Illustration and studies of the family Orchidaceae. The genus Habenaria in North America. Fascicle IV. The Merrymount Press. Arditti, J. et al. 1979. Morphometry of orchid seeds. I. Paphiopedilum and native California and related species of Cypripedium. Am. J. Bot. 66: 11281137. Barthlott, W. and Ziegler, B. 1981. Mikromorphologie der Samenschalen als systematisches Merkmal bei Orchideen. Berichte Deutsch. Bot. Gesells. 94: 267-273. Bateman, R. M. et al. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 2. Infrageneric relationships and reclassification to achieve monophyly of Orchis sensu stricto. Lindleyana 12: 113141. Bateman, R. M. et al. 2003. Molecular phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidadeae). Bot. J. Linn. Soc. 142: 1-40. 82

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Britton, N. L. and Brown, A. 1943. An illustrated flora of the northern United States, Canada and the British possessions. Vol. I. Ophioglossaceae to Polygonaceae. (2nd ed.). New York Bot. Gard. Buttler, K. P. 2001. Taxonomy of Orchidaceae tribus Orchideae, a traditional approach. J. Eur. Orch. 33: 7-32. Case, F. W. 1987. Orchids of the western Great Lakes region. Cranbook Inst. Sci. Chase, M. W. and Pippen, J. S. 1988. Seed morphology in the Oncidiinae and related subtribes (Orchidaceae). Syst. Bot. 13: 313-323. Clifford, H. T. and Smith, W. K. 1969. Seed morphology and classification of Orchidaceae. Phytomorphology 19: 133139. Correll, D. S. 1950. Native orchids of North America north of Mexico. Stanford Univ. Press. Delforge, P. 1994. Guide des Orchidées d‘Europe, d‘Afrique du Nord et du Proche-Orient. Delachaux et Niestle´. Dressler, R. L. 1993. Phylogeny and classification of the orchid family. Timber Press. Gamarra, R. et al. 2007. Seed micromorphology in the genus Neotinea Rchb. f. (Orchidaceae, Orchidinae). Bot. J. Linn. Soc. 153: 133-140. Hapeman, J. R. and Inoue, K. 1997. Plant pollinator interactions and floral radiation in Platanthera (Orchidaceae). In: Givnish, T. J. and Sytsma, K. J. (eds), Molecular evolution and adaptive radiation. Cambridge Univ. Press, pp. 433-454. Healey, P. L. et al. 1980. Morphometry of orchid seeds. III. Native California and related species of Goodyera, Piperia, Platanthera and Spiranthes. Am. J. Bot. 67: 508-518. Kurzweil, H. 1993. Seed morphology in southern African Orchidoideae (Orchidaceae). Plant Syst. Evol. 185: 229247. Löve, A. and Simon, W. 1968. Cytotaxonomical notes on some American orchids. Southwest. Nat. 13: 335-342. Luer, C. 1975. The native orchids of the United States and Canada, excluding Florida. W. S. Cowell Ltd. Mohlenbrock, R. H. 1970. The illustrated flora of Illinois. Flowering plants. Lilies to Orchids. Southern Illinois Univ. Press. Molvray, M. and Kores, P. J. 1995. Character analysis of the seed coat in Spiranthoideae and Orchidoideae, with special reference to the Diurideae (Orchidaceae). Am. J. Bot. 82: 1443-1454. Pridgeon, A. M. et al. 2001. Genera Orchidacearum. Vol. 2, Orchidoideae (part one). Oxford Univ. Press. Rydberg, P. A. 1900. Catalogue of the flora of Montana and the Yellowstone National Park. Mem. New York Bot. Gard. 1: 1-492. Rydberg, P. A. 1901. The American species of Limnorchis and Piperia, north of Mexico. Bull. Torrey Bot. Club 28: 605643. Sheviak, C. J. 1999a. Platanthera hyperborea and a reappraisal of green Platantheras. North Am. Native Orch. J. 5: 117-141. Sheviak, C. J. 1999b. The identities of Platanthera hyperborea and P. huronensis, with the description of a new species from North America. Lindleyana 14: 193-203. Smith, W. R. 1993. Orchids of Minnesota. Univ. of Minnesota Press. Voss, E. G. 1972. Michigan Flora. Part I. Gymnosperms and monocots. Cranbrook Inst. Sci., Univ. of Michigan Herbarium. Webb, D. A. 1980. Platanthera L. C. M. Richard. In: Tutin, T. G. et al. (eds), Flora Europaea 5. Cambridge Univ. Press, pp. 331-332. LITERATURE CITED IN EXTRACTIONS FROM BAUMBACH AND LÜCKEL Bateman, R. M.; Pridgeon, A M. & M. W. Chase (1997); Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 2 - Infrageneric relationships and reclassification to achieve monophyly of Orchis sensu stricto; Lindleyana 12(3);126-127 Britton, Nathaniel Lord, Manual of the Flora of the Northern States and Canada, 1901, Orchidaceae:289-306 Brown, P. M. (2003); The Wild Orchids of North America, north of Mexico. Gainesville: University Press of Florida Brown, P. M. (2008): A new fringed Platanthera (Orchidaceae) from the Central Appalachian Mountains of Eastern North America; North American Native Orchid Journal 14(4):239-254 Brown, P. M. (2008): A long-known, but enigmatic Platanthera hybrid from eastern North America; North American Native Orchid Journal 14(4):255-261 Butzin, F. (1981): Fimbriella, eine neue Orchideenngattung; Wildenowia 11 :321-325 Catling, P. M. & V. Catling (1994): Identification of Platanthera lacera hybrids from New Brunswick and Nova Scotia; Lindleyana 9: 19-32 Correll, D. S. (1950): Native Orchids of North America: 49-116 Farwell, O. A (1901): A catalogue of the flora of Detroit 1901; Reports of the Michigan Academy of Science 2:31-68 Lindley, J. (1835): The Genera and Species of Orchidaceous Plants: 284-288 83

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Luer, C. A. (1972): The native Orchids of Florida: 136-151 Luer, C. A. (1975): The native Orchids of the United States and Canada Excluding Florida :75-93: 145-148 Rafinesque, C. S. (1836): Flora Telluriana, Orchiidaceae part. 2, Cent. 2:38-39 Pridgeon, A. M., Bateman R. M., Cox, A. V., Hapeman, J. R. & M. W. Chase (1997): Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 1 - Intergeneric relationships and polyphyly of Orchis sensu lata; Lindleyana 12(2):89-109 Rydberg, P. A (1901): Family 2. Orchidaceae Lindl. (Orchid Family); Schrenk, W. J. (1976): Die Orchideen der nordosttlichen Vereinigten Staaten, Teil IV; Die Orchidee 27(4):153-158 Schrenk, W. J. (1977a): Die Orchideen der norddostlichen Vereinigten Staaten, Teil VII; Die Orchiidee 28(2):66-69 Schrenk, W. J. (1977b): Zusammenstellung der Orchideenarten der Vereinigten Staaten yon Amerika und der amerikanischen Jungferninnseln; Die Orchidee 28(3):98-104 Senghas. K. (1973): 5. Subtribus: Platanthennae; in Schlechter, R.: Die Orchideen ed.3. ed. IIA (Lieferung 4):202-216 a GLOSSARY allele: one of two or more alternative forms of a gene, occupying the same position (known as a locus) on paired chromosomes and controlling the same inherited characteristics; alternative DNA sequences at the same locus which may or may not result in different phenotypic traits allopolyploidy: the condition of having two or more complete sets of chromosomes derived from different species atavistic: relating to or displaying the recurrence of a genetic feature that has been absent for several generations; the tendency to revert to an ancestral type autogamy: self-pollinations whereby the fertilizing pollen is from the same blossom as is the pistil receptacle basionym: the original previously published legitimate name or epithet from which a new name is suggested for a taxon of different rank or position bursicle: a small sac or pouch-like receptacle clade: a group of organisms consisting of a single common ancestor and all its descendants; a monophyletic group gynostemium: a reproductive structure found in several plant families whereby the male and female parts (stamens and pistil) are fused into a single organ; column heterochrony: changes in the timing or relative rates of development of different tissues within an organism that can be inherited by its offspring which result in changes to size and shape homoplastic: analogous/similar features or characters that are independently derived within separate lineages which are not descended from a recent common ancestor (e.g., octopus eyes and human eyes) indels: heritable insertions or deletions of a segment of DNA in a sequence; can be used as a genetic marker to differentiate among individuals or groups of individuals; if found in coding 84 regions and its length is not a multiple of 3, an indel will cause a frameshift mutation, possibly resulting in the production of a different protein Internal Transcribed Spacer (ITS): refers to a piece of non-functional RNA situated between structural ribosomal RNAs (rRNA) on a common precursor transcript lamella: layers of membranous material often containing polygalacturons (D-galaturonic acid) and neutral carbohydrates (pectins) monophyletic: containing all descendants and their most recent common ancestor in one clade monotypic: a genus with only one species nominate race: the original form or race for which the group was named orthographic: relating to spelling or wording paedomorphosis: phylogenetic change that involves retention of juvenile characters by the adult ; also, a phenotypic and/or genetic change in which the adults of a species retain traits previously seen only in juveniles paraphyletic: a paraphyletic group contains some, but not all, of the descendants from a common ancestor; containing the most recent common ancestor but not all its descendants in one clade parsimonious: very frugal or ungenerous; the most conservative approach with the least number of changes phylogenetic: genetic evolutionary relationships within and between taxonomic levels, particularly the patterns of descent, often branching, from one organism to another plesiomorphic: An ancestral or primitive character useful in defining a monophyletic group polyphyletic: containing members whose most recent common ancestor is not a member of the group (e.g., warm-blooded animals containing mammals and birds but their last common ancestor was not warm-blooded)

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section (as in section of a genus): a genetically or morphologically distinct group of organisms within a specified genus sensu lato (s.l.): in the broadest sense sensu stricto (s.s.): in the narrowest sense sequence (verb): to determine the nucleotide (bases a,c,g,t) composition of a strand of DNA usually limited in length; since DNA is double-stranded, both strands are usually sequenced and compared to each other for accuracy since each base only matches up to one other base on the opposite strand spacer (DNA): The dNA sequence between genes; the term is used particularly for the spacer DNA

between the many tandemly repeated copies of the ribosomal RNA genes subtribe: a taxonomic division of a tribe synapomorphies: apomorphy (derived or specialized character) shared by two or more groups which originated in their last common ancestor synonym: a difference and equally valid scientific name used for a single taxon testa: seed coat developed from the tissue surrounding the ovule trees: a colloquial term used to describe the evolutionary relationships among a group of organisms, much as a family tree is used to document the genealogy of human lineages

Blephariglottis (leucophaea)

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Blephariglottis chapmanii syn.: Platanthera chapmanii

Blephariglottis lacera syn.: Platanthera lacera

Blephariglottis pallida syn.: Platanthera pallida

Blephariglottis praeclara syn.: Platanthera praeclara syn.: Platanthera

Blephariglottis peramoena syn.: Platanthera peramoena

Blephariglottis ×channellii syn.: Platanthera ×channellii

Blephariglottis ×lueri syn.: Platanthera ×lueri

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Blephariglottis ×andrewsii syn.: Platanthera ×andrewsii

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Limnorchis chorisiana syn.: Platanthera chorisiana

Limnorchis aquilonis syn.: Platanthera aquilonis

Limnorchis brevifolia syn.: Platanthera brevifolia

Limnorchis limosa syn.: Platanthera limosa

Limnorchis dilatata Limnorchis yosemitensis var. leucostachys syn.: Platanthera yosemitensis syn.: Platanthera dilatata var. leucostachys 87

Limnorchis holochila (Hawai‘i) syn.: Platanthera holochila

L. hyperborea (Greenland) syn.: Platanthera hyperborea

Rydberg: LIMNORCHIS

THE AMERICAN SPECIES OF 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

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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

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WILD ORCHIDS IN THE 21ST CENTURY

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

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WILD ORCHIDS IN THE 21ST CENTURY

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

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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
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WILD ORCHIDS IN THE 21ST CENTURY

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

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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

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Empiricist :…..OF CABBAGES AND KINGS

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
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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

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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://magazinedirectory.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.
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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
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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.
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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.

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Kieffer: PLATANTHERA ×CANBYI

Platanthera blephariglottis (left) and P. cristata (right)

Platanthera ×canbyi Photos by Chelsea Kieffer

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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

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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

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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.

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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
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or directly from the author at naorchid@aol.com
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