december 2008 north american native orchid journal

NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 14(4) 2008

IN THIS ISSUE: A NEW SPECIES OF FRINGED PLATANTHERA FROM THE CENTRAL

APPALACHIAN MTS. OF EASTERN NORTH AMERICA AN AUDIENCE WITH THE QUEEN A LONG-KNOWN, BUT ENIGMATIC, PLATANTHERA HYBRID FROM EASTERN

NORTH AMERICA ORCHIDS OF ATLANTIC COAST BARRIER ISLANDS FROM NORTH CAROLINA TO

NEW YORK

and more………….

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

NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 14 (4) 2008

CONTENTS

NOTES FROM THE EDITOR 237

A NEW SPECIES OF FRINGED PLATANTHERA FROM THE CENTRAL APPALACHIAN

MOUNTAINS OF EASTERN NORTH AMERICA Paul Martin Brown, Clete Smith & J. Scott Shriver

238 A LONG-KNOWN, BUT ENIGMATIC, PLATANTHERA HYBRID

FROM EASTERN NORTH AMERICA Paul Martin Brown

254 AN AUDIENCE WITH THE QUEEN

Theo Witsell 262

OBSERVATIONS ON HEXALECTRIS NITIDA FROM CENTRAL TEXAS AND A NEW COLOR FORM

Matt White & Paul Martin Brown 267

TWO COLOR FORMS FROM THE CENTRAL APPALACHIANS Paul Martin Brown

270 THE CORRECT GENUS FOR THE JINGLE BELL ORCHID, HARRISELLA PORRECTA

Barbara Carlsward & Mark Whitten 272

ORCHIDS OF ATLANTIC COAST BARRIER ISLANDS FROM NORTH CAROLINA TO NEW YORK

Eric E. Lamont & Richard Stalter 275

THE FUTURE…..? The Slow Empiricist

288

BOOK REVIEWS 290

ORCHIDS OF RUSSIA AND ADJACENT COUNTRIES THE MARIE SELBY BOTANICAL GARDENS

ILLUSTRATED DICTIONARY OF ORCHID GENERA

NEW TAXA AND COMBINATIONS PUBLISHED IN THE NORTH AMERICAN NATIVE ORCHID JOURNAL VOLUME 14, 2008

293

Unless otherwise credited, all drawings in this issue are by Stan Folsom. The opinions expressed in the Journal are those of the authors. Scientific articles may be subject to peer review and

popular articles will be examined for both accuracy and scientific content. Volume 14(4): 237-296 issued October 15, 2008.

Copyright 2008 by the North American Native Orchid Journal Cover: Epidendrum magnoliae by Stan Folsom

NOTES FROM THE EDITOR

This final issue of 2008 is a momentous one as the Journal presents not only a

new species, but a new hybrid and several new color forms, and is please to offer three articles previously published in essentially non-orchid oriented publications. After their many years of observations and collecting data I have been privileged to work with Scott Shriver and Clete Smith in describing Platanthera shriveri, honoring Scott‘s late father, Albert Shriver. This new species is a distinctive and beautiful addition to the orchid flora of North America. Even more time has been spent by many eastern orchid observers puzzling over a possible hybrid between Platanthera psycodes and P. grandiflora. That mystery is now somewhat solved with P. ×enigma. The electronic format continues to be well received and we now reach more than 1200 readers. You may read this and back issues at: http://wiki.terrorchid.org/tow:journals. The current update of the North American Personal Checklist is also available at that website and will continue to be updated quarterly as needed. Paul Martin Brown Editor [email protected] 10896 SW 90th Terrace, Ocala, Florida 34481 PO Box 759, Acton, Maine 04001 June-mid October

As we begin our 15th year of publication

Coming in January 2009

A FAMILY ORCHID VACATION TO THE GREAT LAKES REGION AND POINTS BEYOND

MORPHOLOGICAL VARIATION IN HABENARIA MACROCERATITIS NATIVE ORCHID FLORAS AND PUBLICATIONS —2007-2008

PLATANTHERA HYBRIDS FROM WESTERN NORTH AMERICA TWO NEW FORMS OF THE FLORIDA ADDER’S-MOUTH

and more!!!!!

237

http://wiki.terrorchid.org/tow:journals

mailto:[email protected]

Brown, Smith & Shriver: A NEW FRINGED PLATANTHERA (ORCHIDACEAE) FROM THE CENTRAL APPALACHIAN MOUNTAINS OF EASTERN NORTH AMERICA

238

FIG. 1. PLATANTHERA SHRIVERI P.M. Brown


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A NEW FRINGED PLATANTHERA (ORCHIDACEAE) FROM THE

CENTRAL APPALACHIAN MOUNTAINS OF EASTERN NORTH AMERICA

Paul Martin Brown, Clete Smith & J. Scott Shriver

ABSTRACT

A new species of fringed orchid, Platanthera shriveri, from the central Appalachian Mountains, is described and illustrated. It differs from the similar P. grandiflora (Bigelow) Lindley and P. ×keenanii P.M. Brown in its flowering time, deeply laciniate lip segments, anther position, spur length, and longer persisting floral bracts. Key Words: Orchidaceae, Platanthera, West Virginia, Virginia, North Carolina, Pennsylvania, hybrid orchids

INTRODUCTION

For more than ten years Shriver and Smith, along with Douglas Jolley, William Grafton, Thomas Sampliner, Stanley Bentley, Rodney Bartgis, and others, have been observing a distinctly different purple fringed orchid in the central Appalachians of West Virginia. Several substantial pure colonies were observed that show no evidence of possible hybridization, and flowering time is significantly later than that of the similar Platanthera grandiflora. Often accompanied on these forays by Clete Smith‘s dear friend and Scott Shriver‘s late father, Albert, the authors have described and named the new species in his honor. For many of these years the plants in question were known among those observing them as the ‗pale frilly orchids‘.

Platanthera shriveri P.M. Brown, sp. nov. TYPE: USA, West Virginia, Pocahontas County, along Williams River Rd. (FS Rd. 216); 5.3 miles south of junction of FS 86 and 216. 17 July 1993. J. Scott Shriver 97 with Albert Shriver, Clete Smith. (HOLOTYPE: CM). (Figures 1, 2, 3, 4a, 7a, 8 left, 9c, 10 upper right, 11).

Planta similis Platanthera grandiflora (Bigelow) Lindley, sed segmenta labelli profunde

laciniata, isthmus labelli longior, calcar longior, inflorescentia laxior, labellum concavum, petala erecta marginatis laciniatis, petala non coniuncta sepalo medio, sepalum medium basi angustatum.

Similar to Platanthera grandiflora (Bigelow) Lindley but differing in the deeply laciniate lip segments, longer isthmus of the lip, longer spur, consistently lax inflorescence, concave lip, and erect spreading petals with lacerate margins, free from dorsal sepal, and tapered to the base.

Plants 35-110 cm. tall; leaves 3-4, wide-spreading and recurved to somewhat ascending, scattered along stem, gradually reduced to bracts distally; blade lanceolate, elliptic, oblong-elliptic, or oblong-obovate, the lower ones 5.0-6.5 cm wide × 12-18 cm long, the lowest leaf usually somewhat smaller than the second leaf; inflorescence lax, 6-25 cm tall × 5-6 cm in diameter with (8)15-22(40) clearly separated flowers, resupinate, showy, pale lavender to rose-purple, floral bracts 3-5 mm wide by 30-45 mm long, persisting well beyond anthesis; lateral sepals strongly reflexed, often marked vertically with darker veins; petals ovate, erect, gradually tapered to base, margins clearly lacerate, often exceeding the dorsal sepal in


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length by up to 5 mm, wide-spreading and free from the dorsal sepal, not forming the often traditional ‗Platanthera hood‘ over the column; lip deeply 3-lobed, porrect, projecting forward, concave or strongly ‗cupped‘, without basal thickening, 13–20 mm long × 18–25 mm wide, lobes deeply laciniate throughout; isthmus white, slender, usually (½)¾+ the length of the lip; spur tapered to apex, 21–35 mm long = to ca. 1.75–2 times the length of the lip; rostellum lobes directed forward, spreading, concave; ovary slender to stout 13–28 mm; fruit a capsule, usually 2-10 per raceme, subtended by the persistent, rather showy bracts (Fig. 8) Habitat: in partial to full shade of damp, open, mixed deciduous and coniferous woods, often along seepage springs or streams, or on roadside banks amid mosses, ferns, grasses, sedges, and/or nettles in mountains at elevations between 850-1,450 m (2,350-4,000 feet). Flowering period: mid-July to early August Etymology: shriveri in honor of Albert Shriver, Jr. (1923-2008) Common name: Shriver‘s frilly orchid Paratypes: NORTH CAROLINA: Watauga Co., Rich Mt. July 24, 1975. D.E. Boufford 17620 & E. Wood

(NCU); PENNSYLVANIA: Somerset Co., Buckstown. July 16, 1905. B.H. Patterson s.n. (CM); VIRGINIA: Highland Co., along U.S. 250 north facing slope, 0.2 miles east of West Virginia state line, August 1, 2008. C.

Garratt s.n. photo & D. Jolley (CM, WVA); WEST VIRGINIA: Pocahontas Co., along USFS 14 (Wymer-Thornwood/Middle Mt. Rd.) between 5.1-10 miles s of Wymer. July 29, 1995. J.S. Shriver 409 with A. Shriver & C. Smith (WVA ; CM); along USFS 14 (Wymer-Thornwood/Middle Mt. Rd.) between the head of Fivemile Hollow and Camp Pocahontas, July 29, 1995. J.S. Shriver 409 with A. Shriver and C. Smith, (WVA); Randolph Co., July 30, 1938. E.E. Hutton s.n. (WVA).

As expected, the 1905 Patterson collection at CM was originally identified as Blephariglottis grandiflora and subsequently by E. Wherry as Habenaria fimbriata, Platanthera grandiflora by Stoutamire in 1972, and Shriver f. as P. ×keenanii in 1996. The more recent collections of Shriver et al. were annotated as P. ×keenanii.

An additional interesting specimen collected by the W.V.U. Biological Expedition on July 6, 1949, is also housed at WVA. The plant is in tight bud and Shriver f. comments on the sheet (1996 annotation) that ‗plant only in bud so identity must be based upon expected (ca. July 15-20) probable bloom date, vegetative appearance, and locality of collection….‘ At that time he annotated it as probable Platanthera ×keenanii and the specimen now would be considered probable P. shriveri. This collection had been previously annotated as Habenaria bracteata (1949 as collected), H. viridis 1978 S. Norris and ―may not be H. viridis‖ G. White. DISCUSSION

Plants of an apparently unique purple-flowered Platanthera were first noted in the field in West Virginia by Smith and the Shrivers in 1992. Following the publication of Platanthera ×keenanii in 1993 by Brown, those West Virginia plants were cautiously assigned to that taxon. Noting distinct differences from more typical plants of P. ×keenanii found nearby, the trio continued to search carefully for more sites and Shriver f. meticulously documented each site and the plants therein. They found only one site of the typical P. ×keenanii. It was observed from 1992-1995 and produced only a few flowering plants during that time. Subsequent searches for more plants of the hybrid in the ensuing years have not yielded any plants. Platanthera ×keenanii may be one of the rarest orchids to be found in the central Appalachians.


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× 1

Fig. 2. Platanthera shriveri P.M. Brown Shriver’s frilly orchid


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Platanthera shriveri flowers about three

weeks after P. grandiflora in sympatric areas and is usually still in bud when P. grandiflora is ending its period of bloom (Fig. 5). Stoutamire (1972) states: Platanthera grandiflora has also been collected in flower during the first half of June in the Carolina-Virginia-West Virginia area…..Flowering sequences are more complex in Virginia and in North Carolina where P. grandiflora has been collected and observed to flower in early August….. at 5000-6000 ft altitude, and this disjunct flowering sequence cannot be due to altitudinal differences. More than one phenological race of P. grandiflora may be present in the southern Appalachians, each being adapted to different broods of pollinators.

Following the lead of Stoutamire

cited above, a reexamination of specimens of Platanthera grandiflora from several herbaria with significant Appalachian collections was conducted. Specimens exhibiting late flowering dates, after July 15th, were flagged and examined closely. Several of these late flowering specimens proved to be P. shriveri rather than P. grandiflora (cited in the paratypes). These specimens represent historic P. shriveri occurrences and include a single specimen from both North Carolina and Pennsylvania and three specimens from West Virginia. The geographic range of P. shriveri, as a result of these historic discoveries in conjunction with extant populations, now extends for 300 km and includes four states- Penn., W. Va., Va., and N. C.

Flowers of Platanthera shriveri are comparable in size to those of P. grandiflora with respect to lip length and lip width, but differ noticeably in ovary length, spur length, and differ to an even greater degree in average fringe depth. Plants of P. shriveri possess longer ovaries (ca. 3.5 mm longer), longer spurs (ca. 6.2 mm longer), and deeper fringing (ca. 4.0 mm deeper). Also, the general look of the raceme is different when one compares the ‗huskiest‘ specimens of the

HISTORICAL PERSPECTIVE:

On July 17, 1992 we met Hal Horwitz and

Stan Bentley while on a trip to the Cranberry

Glades area in West Virginia. They directed us

to a Platanthera ×andrewsii site along the Williams

River Road. There were a few plants that were

undoubtedly P. ×andrewsii (later, after 1994, we

realized they were actually P. ×keenanii). This

was our first encounter with ‗P. grandiflora

looking‘ plants.

In the next few years we continued going to

this site but the Platanthera ×keenanii dwindled

down to where we saw nothing that resembled

that first encounter. However, we were noticing

other plants blooming at this time that we

continued calling P. ×keenanii. These plants were

not at all similar to the original hybrids we saw.

They were much ‗showier‘ than the typical, but

beautiful, P. grandiflora. They even bloomed

slightly later than when we had anticipated seeing

the P. ×keenanii. The plants would be in full-

flower to half-flower at this time. Also, plants

flowered not only by the roadside where we

normally would see the P. grandiflora and P.

×keenanii, but also deeper in the woods. We are

sure these plants went unnoticed until we were

fortunate enough to see them on one of our

numerous trips to the area.

In subsequent years we found the other sites

along Forest Road 14 (Wymer-Thornwood

Road). We found the large site at the South end,

but there were numerous smaller sites and single-

plant sites along this 30 mile road. We started to

realize we were seeing something different, more

stable, than the hybrid P. ×keenanii. All the

plants had the definitive upward aspect of the

more frilly lip. We then noticed the much longer

spur and its upward horizontal aspect (fig. 4). It

was at this time we referred to this plant as ‗the

late blooming grandiflora‘. This was July of 2001.

The large Cranberry Glades site was added and

this was when we knew we had a very different,

viable plant and we started referring to this plant

as the ‗pale frilly‘. We now have the addition of

the large Virginia site and we feel that in the next

few years we will add more sites to this list.

Clete Smith


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P. shriveri to the ‗huskiest‘ specimens of P. grandiflora: the P. shriveri racemes display a much more open appearance with fewer flowers allowing for more empty space between them, whereas P. grandiflora racemes are denser, having flowers which overlap to a slight degree, preventing any space between them. In addition, the fringed lip of the P. shriveri seems to display a cup-like or concave shape to a far greater extent than the typical P. grandiflora lip.

Plants of Platanthera shriveri have been observed recently at five extant stations, each

with populations of 25 to 75 plants, in the Appalachian Mountains of eastern West Virginia and Virginia, each with populations of 25-75 individuals. Two sites are located near Cranberry Glades, one site near Thornwood with all three sites in Pocahontas County, West Virginia, and one site consists of widely scattered individuals or small groupings spread along a 10-miles section of a Forest service Road in Randolph County, and the most recent recorded site for the species occurs along a US Highway west of Monterey, Virginia. The Virginia site, discovered by Charles Garratt, of Warm Springs, Va., and Doug Jolley, of Heaters, W. Va., on August 1, 2008 is especially significant because it represents a new state record for P. shriveri (Fig. 6)!

It should be noted that in an effort to understand the identity of the purple fringed

Platanthera species in the Appalachians, one must ascertain whether a given population represents a stand of a single species or perhaps a hybrid grouping that might involve natural crossing between any of several native species. Making such a determination is not an easy task. Most populations of purple fringed orchids studied in the Appalachian Highlands by Clete Smith, J. Scott Shriver, and others who were involved in the elucidation of P. shriveri have been found to exist in completely pure stands of a single species. When in pure stands, Platanthera grandiflora is by far the most common of the purple fringed orchids at high elevation in the Appalachians while P. shriveri is less common. Platanthera grandiflora and P. shriveri are easy to separate as distinct species based upon morphology and flowering date. There is, however, one notable population that has caused considerable confusion in the separation of P. shriveri from P. grandiflora over the last 15 years. In Pocahontas County, West Virginia, along the nine-mile length of a US Forest Service Road near Cranberry Glades, as many as 250 purple-colored Platanthera orchids of questionable identity can be observed apparently representing both taxa. Along this road both P. shriveri and P. grandiflora are present, at least from the standpoint of morphology, and in many cases plants which appear to represent a melding of the two species, putative hybrids, may be seen. Flowering dates, as a means of separating P. shriveri from P. grandiflora, break down in this population as some individuals which appear morphologically to favor P. grandiflora can be found in flower after July 5th (after P. grandiflora is typically finished) while plants which appear morphologically to favor P. shriveri can be found in bloom prior to July 8th (before P. shriveri typically commences flowering). Intermediate plants may represent the possible hybrids between P. grandiflora and P. shriveri. To further complicate the identity of the various purple Platanthera along this road, the very distinctive P. × keenanii has been found here as well.

[ Note: A similar situation exists in the Osceola National Forest in northern Florida where Platanthera

chapmanii (Small) Luer is the dominant species and P. cristata (Michaux) Lindley, flowering a bit earlier, can also be found in small local stands as well as P. ciliaris (Linnaeus) Lindley. Hybrids between P. ciliaris and P. cristata, P. ×channellii Folsom, are occasionally found as well as the occasional hybrid between P. chapmanii and P. cristata, P. ×apalachicola P.M. Brown & S.L. Stewart, each maintaining their own taxonomic identity (Brown and Stewart, 2003).]


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The geographic distribution of several Platanthera species native to Appalachia is significant if one expects to understand P. shriveri as a distinct taxon. Platanthera grandiflora is rather frequent and occurs in 34 counties of both Appalachian Pennsylvania and West Virginia. Platanthera lacera has an even greater distribution and may be seen in more than 50 counties of Appalachian Pennsylvania and West Virginia. Platanthera shriveri, however, seems to be a far less common species. Platanthera shriveri is an Appalachian endemic, as far as is currently known, and occurs at fewer than a half dozen extant sites scattered in two West Virginia counties and in a single county of Virginia. Herbarium studies have uncovered a couple of historic records for P. shriveri which extends the range of the species to the Appalachian Highlands of both Pennsylvania and North Carolina. At the Carnegie Museum Herbarium in Pittsburgh, a voucher for P. shriveri indicates the historical presence of the species in Somerset County, Penn. This P. shriveri specimen was collected near Buckstown by B. H. Patterson on July 16, 1905. Another voucher for P. shriveri at the North Carolina University Herbarium indicates the species once grew in North Carolina. Platanthera shriveri was collected from Rich Mountain in Watauga County. This P. shriveri specimen, collected by D.E. Boufford and E. Wood on July 24, 1975, represents a significant southern range extension for the species. Because Shriver‘s frilly orchid has just been taxonomically isolated and revealed to the orchid community, it is quite possible that additional populations may be found throughout the Appalachian Mountains in the next few years as more people seek to locate the species. Perhaps some populations may even be located beyond the boundaries of Appalachia. The presence of an orchid endemic to the Appalachian region is not unique as Corallorhiza bentleyi Freudenstein, with a similar restricted distribution in West Virginia and Virginia was described in 1999. Another orchid, Listera smallii Wiegand, and numerous other non-orchids are also known as endemics or near-endemics to the region.

Interestingly, the hybrid Platanthera ×keenanii, in the Appalachians may actually be the

rarest of all the Platanthera taxa as it is known from only a single population in West Virginia. This population has waned over the years and plants fitting the description of P. ×keenanii are no longer extant anywhere in the region. Perhaps P. ×keenanii, as is the case with many hybrids, represents a rather unstable and short-lived taxon. Keenan's hybrid orchid should continue to be sought throughout Appalachia, especially in areas where its parents bloom together.

Taxon Relationships The section Blephariglottis of Platanthera, found in North America, presents several

of the showiest native orchids found on the continent. Although there have been only three purple-flowered species–P. peramoena (A. Gray) A. Gray, P. psycodes (Linnaeus) Lindley, and P. grandiflora–recognized for many years, a fourth member of this section with greenish-white-yellowish flowers–P. lacera (Michaux) G. Don–is an important component for it has contributed to two distinctive hybrids: P. ×andrewsii (Niles) Luer (P. lacera × P. psycodes) and P. ×keenanii (P. lacera × P. grandiflora). A fourth purple-flowered species is presented here–P. shriveri.

Platanthera shriveri presents a situation that is not unique among North American

Platanthera species. Two taxa are known from southeastern United States–P. chapmanii (Small)


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Luer and P. ×channellii Folsom–bearing strong superficial resemblances but having subtle morphological differences (Folsom, 1984). Folsom‘s work set a precedent for recognizing both the contemporary hybrid–P. ×channellii, and the species–P. chapmanii. The major difference is that P. chapmanii is a stable, reproducing species but with a very limited distribution in the panhandle of Florida, historically in adjacent Georgia, and in eastern Texas (Brown, 2007) and P. ×channellii is an infrequent, if not rare, randomly occurring hybrid throughout much of the range of its two parents, P. ciliaris and P. cristata. Because P. chapmanii is intermediate in morphology, it has been hypothesized that that same parentage, P. ciliaris and P. cristata, gave rise to P. chapmanii (Folsom, 1984; Brown and Folsom, 2007). Platanthera pallida P.M. Brown from eastern Long Island may also represent this same scenario with the ancestral parentage being P. blephariglottis (Willdenow) Lindley and P. cristata (Michaux) Lindley and the contemporary hybrid being P. ×canbyi (Ames) Luer. Again, there is a superficial resemblance but very distinctive differences and P. pallida grows in a few restricted pure stands of several hundred plants (Brown, 1992, 2008) (Fig. 10).

In determining the status of Platanthera shriveri it is not so much a question of how

these plants differ from P. grandiflora, but how they differ from P. ×keenanii. Chart 1 shows comparative data for those two taxa and applicable data for P. ×keenanii. In the case at hand two superficially similar taxa, P. ×keenanii and P. shriveri, again having subtle morphological differences, are found within the range of the putative parents of the hybrid taxon. In situations such as this, one of the best empirical tests to determine the possibility of which taxon is present is to take note of where they are growing and if they are sympatric with other Platanthera species. Hybrids almost always occur in small numbers or even as individuals with both parents nearby (although given the lifespan of some individuals only one parent may be immediately present); whereas a species usually will occur with multiple individuals in pure stands and/or with other species of Platanthera nearby. The aforementioned hybrids tend to set little seed and rely upon active hybridization of the parents to produce more plants. Only in Nova Scotia, New Brunswick and, especially, in western Newfoundland do P. lacera and P. grandiflora (and P. psycodes) occur in such numbers and flower simultaneously, enable the creation of hybrid swarms with multitudes of apparent backcrosses. There it is often difficult to find pure individuals of any of the three parent species (Brown and Folsom, 2006; Catling and Catling, 1994)! In West Virginia this is neither the case, nor the problem.

Populations of Platanthera shriveri in Appalachia are well defined and are known to

reproduce in pure stands in every extant station. Two of these pure stands have been closely monitored and examined for 15 years and consist of 25-75 individuals, each with about an equal distribution of mature, flowering plants and juvenile, seedling plants. Platanthera shriveri at these sites demonstrates a high degree of morphological floral uniformity, and any of the variations noted in individuals tend to be minor and are associated with attributes like plant height, purple color intensity (pale vs. deep), or vertical veining, which may or may not be well visible on the dorsal sepal of some plants. Uniformity is not only evident within a given population in any year, but also remains true over several years, at least as is indicated by the past 15 years of observations. Also, P. shriveri populations separated by distances of up to 50 miles are also florally identical, so the species shows virtual uniformity throughout its entire range. As an implication of these observations, it is hypothesized here that P. shriveri


246

represents a distinct, long-lived, modern taxonomic species of possible ancient hybrid origin with P. grandiflora and P. lacera as parents. Platanthera shriveri superficially resembles, but differs significantly, from the local, occasionally observed, short-lived contemporary hybrid P. ×keenanii, with the same putative parentage, which is quite rare in Appalachia.

Sharing a similar scenario regarding origin, the three species considered here to be of

ancient hybrid lineage: Platanthera shriveri, P. chapmanii, and P. pallida, are all restricted in their distribution, but the ancestral parents are widespread and relatively common. These same three species occur at or near the edge of the ranges of the parents and individuals of the respective species are reliably uniform in their morphology. In the case of all three contemporary hybrid taxa (P. ×keenanii, P. ×canbyi, and P. ×channellii), a certain amount of morphological variability is found among individuals that exhibit the various traits of the parents.

Fig. 3 Platanthera shriveri illustrating the

distinctive lip and spur ×2


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KEY TO THE PURPLE-FLOWERED PLATANTHERA SPECIES (see figure 7) 1a spur orifice a transverse oval, hourglass on its side, or a transverse dumbbell…P. psycodes

1b spur orifice rounded, rectangular, or angled…2

2a lip margin denticulate to very shallowly fringed…P. peramoena

2b lip margin fringed to more than ⅓ the depth of the lip segments…3

3a lip segments moderately to deeply fringed; isthmus stout, length ca. twice the width; spur 1¼ times

the length of the lip; orifice round….P. grandiflora

3b lip segments deeply and compoundly lacerate; isthmus slender, length ca. 4 times the width; spur 2-2½

times the length of the lip; orifice angled at top…..P. shriveri

Accommodating a hybrid in the above key is very difficult and also very artificial as plants of P. ×keenanii may

easily look like purple-flowered P. lacera, green- or white-flowered P. grandiflora with slender, lacerate lip

segments or any of the above combinations! Platanthera shriveri, on the other hand, does not show this variability

at all.

× ½

a: Platanthera shriveri b: Platanthera ×keenanii c: Platanthera grandiflora

× 2

Fig. 4. Comparison of individual flowers


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10││││15││││20││││25││││30││││5││││10││││15││││20││││25││││30

June July P. grandiflora 2007 2008 P. shriveri 2007 2008 Prime flowering dates 6/23/07 6/27/08 7/14/07 7/18/08

Fig. 5 Prime flowering time separation of Platanthera grandiflora and P. shriveri

Fig. 6.

Distribution

of Platanthera shriveri

●= extant sites ○ = historical sites

○

●

● ●

●

PA

Somerset

Randolph

Pocahontas Highland

VA

WV

●

○

Watauga NC


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P. shriveri P. grandiflora P. ×keenanii inflorescence habit loose; flowers compact; flowers loose well spaced approximate lateral sepals strongly reflexed forward to reflexed reflexed petals spatulate, gradually ovate-oblong abruptly slender, spatulate,

tapering to base tapering to base erect erect and spreading overlapping with

dorsal sepal

lip deeply and com- incised usually to ca. slender lobes deeply poundly laciniate ⅓+ the lip length and irregularly cut

isthmus narrow; = to ¾ broad; = to ⅓ the slender

the lip length lip length spur tapering to apex, clavate ca.1½× length slender, clavate = to

1¾-2 × length of lip of lip ¾+ × lip length orifice tapered at both ends rounded rounded rostellum lobes widely spreading, spreading, convex parallel concave

Chart 1. Comparisons of morphological characters

a: P. shriveri b: P. grandiflora c: P. ×keenanii Fig. 7. Column and

orifice ×5 d: P. lacera


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Fig. 8. Infructescences of

Platanthera shriveri (left) and

P. grandiflora (right) × 1

a: P. grandiflora horizontal, clavate spur round orifice simple fringing on margin of lip


b: P. lacera

descending, essentially clavate spur

erect petals

sparsely bifid, slender lip segments

b: P. lacera descending, essentially clavate spur erect petals sparsely bifid, slender lip segments

d: P. ×keenanii erect to spreading, non incised petals rounded orifice irregularly bifid, incised slender lip segments

Fig. 9. Individual flowers ×3

c: P. shriveri long, tapered horizontal to somewhat ascending spur spreading, prominently incised petals compound bifid, deeply fringed, flabellate lip segments


d: P. ×keenanii erect to spreading, non incised petals rounded orifice irregularly bifid, incised slender lip segments

Fig. 9. Individual flowers ×3


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P. grandiflora P. ×keenanii P. lacera P. shriveri

P. cristata P. ×canbyi P. blephariglottis P. pallida

P. cristata P. ×channellii P. ciliaris P. chapmanii

Fig. 10. Comparisons of three (ancient hybrid) species (right column), possible parentage, and contemporary hybrids (left column) ca. ×⅓


252

LITERATURE CITED: Brown, P.M. 1992. Platanthera pallida (Orchidaceae), a new species of

fringed orchis from Long Island, New York, U.S.A. Novon 2: 308-

11.

______. 2002. Revalidation of Platanthera conspicua. North American

Native Orchid Journal 8: 3-14.

______. 2008. Platanthera pallida - fifteen years of comparisons. North

American Native Orchid Journal 14(2): 157-66.

Brown, P.M. and S.N. Folsom. 1993. A Field and Study Guide to the

Orchids of New England and New York. Jamaica Plain, Mass.: Orchis

Press.

______. 2006. Wild Orchids of the Canadian Maritimes and Northern Great

Lakes Region. Gainesville: University Press of Florida.

Brown, P.M. and S.L. Stewart. 2003. Two new Platanthera hybrids.

North American Native Orchid Journal 9: 35.

Catling, P.M. and V. Catling. 1994. Identification of Platanthera lacera

hybrids from New Brunswick and Nova Scotia. Lindleyana 9: 19-32.

Freudenstein, J. V. 1999. A new species of Corallorhiza (Orchidaceae)

from West Virginia, U.S.A. Novon 9: 511–13.

Folsom, J. P. 1984. Una reinterpretación del estatus y relaciones de las

taxa del complejo de Platanthera ciliaris. (A reinterpretation of the

status and relationships of taxa of the yellow-fringed orchid

complex.) Orquidea (Mexico City) 9: 321–45.

Stoutamire, W.P. 1974. Relationships of purple fringed orchids

Platanthera psycodes and P. grandiflora. Brittonia 26: 42-58.

ACKNOWLEDGEMENTS: Photographs of Platanthera shriveri, P. ×keenanii, P. grandiflora, P. lacera,

and Albert Shriver by J. Scott Shriver excepting figs. 3 & 4a, by Clete

Smith and fig. 11 by Shirley Curtis

Photographs of Platanthera cristata, P. ciliaris, P. chapmanii, P. pallida, P.

×canbyi, P. ×channellii by Paul Martin Brown

Line art:: fig. 2 by Stan Folsom, fig. 9, by J. Scott Shriver;.

The authors thank Bonnie Isaac, Collection Manager, Carnegie

Museum Herbarium (CM), Donna Ford-Werntz, Collection Manager,

West Virginia University Herbarium (WVA) for the loan of specimens

and Alan Weakly and the staff of University of North Carolina

Herbarium (NCU) for facilitating digital specimen scans, and Conley K.

McMullen, James Madison University (JMUH); the managers and/or

staff of NY, HUH, TENN, CLEM, PA, WCUH, DUKE, ASUH for

access to databases and checking specimens. Stan Folsom, Helen Jeude,

Jim Fowler, Scott Stewart, Tom Nelson, Dietrich and Ursula

Rueckbrodt, Sally Puth, Shirley Curtis, and two anonymous reviewers

made several helpful suggestions.

Paul Martin Brown, Research Associate, University of Florida

Herbarium, Florida Museum of Natural History, 379 Dickinson Hall,

PO Box 110575, Gainesville, FL 32611-0575 [email protected]

Clete Smith, 332 Fieldbrook Dr., Pittsburgh, PA 15228 [email protected]

J. Scott Shriver, 110 Hillvue Lane, Pittsburgh, PA 15237

[email protected] Fig. 11. Platanthera shriveri


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Fig. 12. Albert Shriver with one of his precious orchids (1995).

Brown: A LONG-KNOWN, BUT ENIGMATIC, PLATANTHERA HYBRID FROM EASTERN NORTH AMERICA

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Fig. 1 Platanthera ×enigma P.M. Brown Cornish, Maine 30 July 2008


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A LONG-KNOWN, BUT ENIGMATIC, PLATANTHERA HYBRID FROM EASTERN NORTH AMERICA

Paul Martin Brown

Natural hybrids in the genus Platanthera in North America are not at all unusual and in

colonies of two or more species such hybrids are often detected. Hybrids among the fringed-lipped Platanthera species are certainly not novel and nearly every combination possible has been published or noted (Brown, 2008; Brown and Folsom, 2002). Eleven hybrid taxa or nothospecies have been published in the fringed-lipped section of Platanthera (Appendix 1.) The following may now be added to the list. Platanthera ×enigma P.M. Brown nothsp. nov. TYPE: U.S.A. Maine: York Co. wet (usually standing water) roadside ditch along state highway 5 south of Cornish, with graminoids and Ilex verticillata bordered by Tsuga canadensis. 3 August 2008. P.M. Brown 30808 (holotype: MAINE). (Figures 1, 2, 3b, 4b, 5). Planta inter Platanthera grandiflora (Bigelow) Lindley et P. psycodes (Linnaeus) Lindley intermedia et habitu, colore et forma florum, vel proprietibus speciearum mixtis Intermediate in characters between the two parents Platanthera grandiflora and P. psycodes ETYMOLOGY: enigma from the Latin meaning something obscure, hard to understand, or explain

Platanthera grandiflora (Linnaeus) Lindley (figs. 3a, 4a) and P. psycodes (Linnaeus) Lindley (figs.

3c, 4c) often grow in similar moist habitats and even in the same physical areas in northeastern North America. For much of the range, especially in the southern half of this area, the species do not have an overlap of flowering times. But in northern New England, the Canadian Maritimes, and especially southwestern Newfoundland both species may be frequently found flowering together (Stoutamire, 1974). To complicate this scenario P. lacera (Michaux) G. Don is also often found with them. This has resulted in hybrid swarms in Nova Scotia, Prince Edward Island, and Newfoundland that often defy specific parentage as backcrosses are prevalent (Catling and Catling, 1997). But in much of New England, although the two species also grow with P. lacera, the flowering times are more markedly separated. Only in far northern Vermont and New Hampshire does one find locales with both purple species and occasional P. lacera flowering sympatrically. In northern York County, southwestern Maine, plants of P. grandiflora flower from late June, in an early year, to late July, with peak flowering in mid July. Platanthera psycodes flowers primarily in August with only a few early plants to be found in late July. Although P. lacera is present most all of this time its habitat is usually quite different—higher and drier—from either of the purple-flowered species and individuals of both the hybrids, P. ×keenanii P.M. Brown and P. ×andrewsii (White in Niles) Luer, are very rare.

Stoutamire (1974) clearly demonstrated the distinctiveness of the two species. Popular works

for many years used the depth of the fringing on the lip as an aid to identification. After examining


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several thousand plants of both species over the years it has become apparent that the relative depth of the fringe in relation to the size of the lip is not a reliable character. In most cases it works well with the fringe being ⅓ or less in Platanthera psycodes and greater than ⅓ in P. grandiflora, but many plants exist of both species that do not conform to these criteria. The only reliable criteria are those of the orifice shape, spur/ovary length, and overall flower size. Flower color is highly variable and in one meadow in Newfoundland that contained both species and P. lacera all color and growth forms could be found as well as a plethora of hybrids and backcrosses!

For over twenty years a roadside ditch near Cornish, Maine has regularly provided flowering

plants (3-12) of a somewhat enigmatic purple-flowered Platanthera. These plants flower with regularity at the end of July and beginning of August. Although typical P. grandiflora may be found not far away it usually flowers in early July and P. psycodes down the road in mid August. Climate being what it may in Maine there have been years when both species have had a brief overlap in flowering time – thus the resulting putative hybrid plants in the ditch. Although hybrids of P. grandiflora and P. psycodes have often been mentioned in the literature and observed in the field, sorting out pure hybrids from the swarms has proven to be less than reliable (Catling and Catling, 1997; Correll, 1939; Fernald, 1950; Sheviak, 2002). For this reason several workers with these species have refrained from formally describing the hybrid.

Here in Cornish the hybrid plants are clearly evident with no indication of backcrossing or

influence from Platanthera lacera. In many ways these are classic hybrids showing the influence of both parents. Overall the plants appear at first glance to be a robust P. psycodes until one looks closely at the flowers and notes the floral size, just over 1 cm across, deeply fringed lip segments, and the more rounded orifice typical of P. grandiflora. The spur orifice is neither the neat circular shape of P. grandiflora nor the distinctive transverse oval or dumbbell of P. psycodes, but more of a rounded square. A note here on the ‗transverse dumbbell‘ shape is in order. This shape is created by a thickened ridge at the roof of the nectary and may or may not be clearly evident in all individuals. If this ridge extending is at the opening of the nectary then the dumbbell shape is created but if the ridge starts just inside the opening then it is less evident and the opening appears more oval in shape, still dramatically different from that of P. grandiflora. Stoutamire (1974) states ―The nectary opening [of P. psycodes] is oblong in face view and often is partially constricted by a projection from the roof of the nectary at the entrance or just inside the orifice, effectively producing two lateral openings, one beneath each viscidium… The degree of constriction of the nectary varies from plant to plant and perhaps with flower age but is distinct from the unobstructed circular opening characteristic of P. grandiflora‖.

Plants of Platanthera ×enigma have been casually identified in the past as small, late ‗grandiflora‘

or early large ‗psycodes‘—both perfectly logical conclusions and suited to the hybrids. One of the most curious things about these plants is that the spur and ovary are heavily tinted with purple, unlike plants observed of P. grandiflora and P. psycodes in this area, although P. psycodes from other geographic areas show this pigmentation (Saulys, 2005).

Chart 1 shows comparative dimensions and measurements of the two parents and the hybrid and figures 3 and 4 show photographs of the three taxa. Light, moisture, and exposure greatly influence the habit of most Platanthera species and to ensure an equitable comparison plants were chosen from similar wet habitats with similar light conditions.


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Fig.2 Platanthera ×enigma Cornish, Maine 30 July 2008


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Fig. 3 a: Platanthera grandiflora b: P. ×enigma c: P. psycodes

_____ = 1 cm

Fig. 4 a: Platanthera grandiflora b: P. ×enigma c: P. psycodes

__________ = 1 cm


259

Fig. 5. The granddaddy of all Platanthera ×enigma This individual was 112 cm tall with stem diameter near base of 2 cm (base of plant not visible as it is in the water) and

the inflorescence was 30 cm with over 200 flowers and buds! Cornish, Maine 3 August 2008


260

P. grandiflora P. ×enigma P. psycodes raceme diameter 4–6 cm 3–4 cm 2–3 cm diameter/height ratio 1:1–1:3 1:3–1:5 1:4–1:8 lip width 2 cm 1.5 cm 1 cm fringe depth 3–10 mm 2–7 mm 1–3 mm spur length (22) 26 (30) mm 20 (22) mm (12) 15 (18) mm ovary length 25 mm 12–18 mm 15 mm orifice shape rounded rounded to oval transverse oval/dumbbell anther placement parallel somewhat angled angled floral bract length 27 mm lower–15 mm upper 30 mm lower–10 mm upper 17 mm lower–5 mm upper

Chart 1. Comparison of morphological characters from 10 individuals from southern Maine

Summary of chart 1: P. grandiflora has proportionately stout, rectangular racemes with eventually all flowers open simultaneously P. psycodes has proportionately slender, elongated, often tapering, racemes with the lower flowers senescing by the time the uppermost flowers open Height of plants (P. psycodes is often much taller than P. grandiflora) and number, placement and/or size of leaves is not pertinent and can vary considerably with both species depending on latitude, altitude, and habitat. In theory plants of P. ×enigma could favor either parent and appear to be small-flowered P. grandiflora or large flowered P. psycodes. In the Cornish site the overall aspect is that of a large, robust P. psycodes. Plants seen in Newfoundland appear to favor P. grandiflora in overall aspect but with small flowers.

Appendix 1. Published hybrids in the fringed-lipped section of Platanthera Platanthera ×andrewsii (White in Niles) Luer (P. lacera × P. psycodes) Bog-trotting for Orchids. P. 258. 1904. Platanthera ×apalachicola P.M. Brown & S.L. Stewart (P. chapmanii × P. cristata) North American Native Orchid Journal 9: 35. 2003. Platanthera ×beckneri P.M. Brown (P. conspicua × P. cristata) North American Native Orchid Journal 8: 3-14. 2002. Platanthera ×bicolor (Rafinesque) Luer (P. blephariglottis × P. ciliaris) Flora Telleuriana 2: 39. 1837 Platanthera ×canbyi (Ames) Luer (P. blephariglottis × P. cristata) Rhodora 10: 70. 1908. Platanthera ×channellii Folsom (P. ciliaris × P. cristata) Orquidea (Mex.) 9(2): 334. 1984.

Platanthera ×enigma P.M. Brown (P. grandiflora × P. psycodes) North American Native Orchid Journal 14(4): 255 Platanthera ×hollandiae Catling & Brownell (P. leucophaea × P. lacera) Canadian Journal of Botany 77(8): 1144-49. 1999. Platanthera ×keenanii P.M. Brown (P. grandiflora × P. lacera) Field and Study Guide to the Orchids of the New England and New York p. 189. 1993. Platanthera ×lueri P.M. Brown (P. conspicua × P. ciliaris) North American Native Orchid Journal 8: 3-14. 2002. Platanthera ×osceola P.M. Brown & S.L. Stewart (P. chapmanii × P. ciliaris) North American Native Orchid Journal 9: 35. 2003. Platanthera ×reznicekii Catling, Brownell & G. Allen (P. leucophaea × P. psycodes) Lindleyana 14(2): 77-86. 1999.

LITERATURE CITED: Brown P.M. 2008. Personal Checklist of the Orchids of North America north of Mexico. Special Publication #4 of the

North American Native Orchid Journal.


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Brown, P.M. and S.N. Folsom. 2002. The Wild Orchids of North America north of Mexico. Gainesville: University Press of Florida.

_____. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region. Gainesville: University Press of Florida.

_____. 2007. Wild Orchids of the Northeast. Gainesville: University Press of Florida. Catling P.M. and V. Catling. 1994. Identification of Platanthera lacera hybrids from New Brunswick and Nova Scotia.

Lindleyana 9: 19-32. Correll, D. S. 1939. A new status for × Habenaria andrewsii. Botanical Museum Leaflet 7(4): 57-68. _____. 1950. Native Orchids of North America North of Mexico. Waltham, Mass.: Chronica Botanica Fernald, M.L. 1950. Gray’s Manual of Botany 8th edition. New York: American Book Company. Saulys, E.S. 2005 Platanthera psycodes images in Connecticut Botanical Society website http://ct-botanical-

society.org/galleries/platantherapsyc.html Sheviak, C.S. 2002. Platanthera pp. 551-71. In: Flora of North America Editorial Committee, eds., Flora of North America

north of Mexico vol. 26. Oxford and New York: Oxford University Press. Stoutamire, W.P. 1974. Relationships of purple fringed orchids Platanthera psycodes and P. grandiflora. Brittonia 26: 42-58.

Acknowledgements: The author thanks Chuck Sheviak and Paul Catling for early discussions concerning the status of this hybrid combination , Stan Folsom for initially taking PMB to the location in Cornish, Maine in 1986 and subsequent sightings through 2008, and Scott Stewart, Jim Fowler, Sylvain Beausejour, Sally Puth, and Shirley Curtis for helpful corrections and comments.

Witsell: AN AUDIENCE WITH THE QUEEN

262

AN AUDIENCE WITH THE QUEEN

Theo Witsell Photographs by Craig Fraiser

Reprinted with permission from Claytonia (Arkansas Native Plant Society Journal) Fall-Winter 2008.

Craig ―Coondog‖ Fraiser and I recently spent two long hot days in the Springfield

Plateau section of the Ozarks exploring sinkhole ponds and two anonymous spring-fed stream gorges*. We had hoped to find some new species for Arkansas – Virginia sneezeweed (Helenium virginicum), forked aster (Eurybia furcata) and tall larkspur (Delphinium exaltatum), but struck out on all counts. But deep down, unspoken even, we were secretly both hoping to discover a new population of the showy lady‘s-slipper orchid (Cypripedium reginae), far and away the rarest and most seldom-seen lady‘s-slipper in Arkansas. It is so rare, and so spectacular, that it often goes by its other name: simply ―The Queen‖.

The rugged limestone gorges of the Springfield Plateau provide habitat for many rare species.

As we hiked up the rugged canyon of our first stream, we searched likely habitat for


263

forked aster (bases of bluffs and limestone ledges with an accumulation of rich, moist soil), but to no avail. But the scenery was spectacular and the water was among the cleanest and clearest I‘ve ever seen in Arkansas. So clear, in fact, that the clarity was deceiving, turning what looked like a knee-deep step into a cold, take-your-breath-away belly-deep plunge. Rare and uncommon plants abounded along the stream, with cascades of running strawberry bush (Euonymus obovata) spilling from blufftops, and sheer walls peppered with the grey-green foliage of the littleflower alumroot (Heuchera parviflora var. puberula). The rock ledges along the stream banks were loaded with plants that indicated the presence of groundwater seepage: golden ragwort (Packera aurea), umbrella sedge (Fuirena simplex var. simplex), shining coneflower (Rudbeckia fulgida), bishop‘s cap (Mitella diphylla), grass-of-Parnassus (Parnassia grandifolia), and bristly-stalked sedge (Carex leptalea). In the woods along the stream we also found ―new‖ populations of several rare species including satin brome (Bromus nottowayanus), blue cohosh (Caulophyllum thalictroides), and butternut (Juglans cinerea). Even the sand grape (Vitis rupestris), by far the rarest of our native grapes, with its wide, folded leaves, was found growing in the gravel of the stream bed.

The hanging garden, high above the stream, keeping the Queen safe from her enemies.

The streams we were traveling in were deeply incised, forming dramatic ‗box canyons‘with bluff walls and narrow floodplains, making walking in the stream channel the most convenient, and


264

perhaps the safest, avenue for travel. Signs of roaring spring floods (drifts of dead leaves crammed head high in the branches of shrub thickets, occasional logjams against trees high above the water, and high, steep-walled gravel bars up against deep, scoured, bedrock-bottomed pools) spoke to the fact that the streams, while they are lazy and docile in the summer, have an excitable mean streak during the wet season. All along, as we waded in the cool water, we scanned the bluffs for the telltale signs of seepage, which would tell us to be on the lookout.

To get to our second stream, we descended over 300 feet down into its gorge from an adjacent ridgetop (there are no roads that cross this particular stream due to the rugged and inaccessible terrain). When we reached the valley floor we arbitrarily decided to go upstream. The water was cold and still running in early August, indicating that springs supply a good portion of the flow. After about a mile of slipping and sliding up the creek, necks craned to survey the bluffs, I did a double-take. There, 16 feet up on a sheer limestone wall, was a small, lush ―hanging garden‖, perhaps five feet wide and three or four feet front-to-back, obviously kept moist by the gentle emergence of groundwater. Against the wall of the bluff, at the back of this secret garden, were ten of the largest lady‘s-slipper orchids I‘ve ever seen! The biggest were perhaps three feet tall, with leaves eight or ten inches in length. The specific epithet, reginae (the Queen), is aptly given.

Surveying the bluffs is best done from the stream channel, but watch out for leeches, be sure to keep your pack out of the water, and always take a waterproof bag for your camera and phone! You‘ll fall in eventually.

Believe me, I speak from experience.


265

The unique and fragile microhabitat supporting this small population was remarkable in itself, as was the assemblage of associate species present – the Queen‘s court. There were a few small, arching shrubs of ninebark (Physocarpus opulifolius) and the uncommon alternate-leaved dogwood (Cornus alternifolia). And there were a number of seepage-loving herbaceous plants: spotted cowbane (Oxypolis rigidior), grass-of-Parnassus, bristly-stalked sedge, shining coneflower, bearded shorthusk grass (Brachyelytrum erectum), and even eastern columbine (Aquilegia canadensis). Everything must have been just perfect… just the right amount of light, the required mycorrhizal fungi to assist the orchids in obtaining nutrients, just the right amount of water, and the inaccessible site itself… a fortified castle to protect the Queen from her enemies, like hungry deer and greedy poachers.

The Queen and her court.

The day, which was hot and humid with thunder clapping in the distance, suddenly seemed brighter with our discovery and I felt somewhat lighter as we slogged back down the creek and up the steep slopes out of the gorge. We didn‘t find what we had really set out to find, but we got a number of nice surprises along the way and got a rare audience with The Queen, something I think we‘ll both remember for a long, long time.


266

Showy lady‘s-slipper orchid (Cypripedium reginae). Photo by John Pelton.

* The names of these streams are not disclosed here because of the unfortunate and continued poaching of lady‘s-slipper orchids, especially the Queen slipper, from the wild by collectors. Several historical populations in Arkansas are now gone because all the plants were dug out. Carl Hunter once told me that, at one time, he knew of five sites for the species in Benton County, but that all of them had been lost to poachers. Today no populations are known to survive in the northwestern part of Arkansas, and only four are known in the entire state. Theo Witsell, Arkansas Natural Heritage Commission, 1500 Tower Building, 323 Center St., Little Rock , AR 72201 [email protected]

White & Brown: OBSERVATIONS ON HEXALECTRIS NITIDA FROM CENTRAL TEXAS AND A NEW COLOR FORM

267

OBSERVATIONS ON HEXALECTRIS NITIDA FROM CENTRAL TEXAS AND A NEW COLOR FORM

Matt White & Paul Martin Brown Hexalectris nitida L.O.

Williams occurs in both cleistogamous and chasmo-gamous flowering forms. The species is known from widespread locales in Texas, a single historical site in New Mexico, and scattered sites in northern Mexico. For several years pale-colored plants of the more common cleisto-gamous form have been observed in at least three counties in Texas (Fig. 4). White‘s observations in Dallas County have proven valuable in understanding this color form. Unlike many ‗colorless‘ forms, lacking in all anthocyanins—the red, purple and/or blue color-ations, the lip in this case is still highly colored as in the typical plants. Figure 3 shows the lip teased open to reveal the coloration. Also growing nearby is the pale form of the Texas H. warnockii Ames & Correll forma flavida Catling (Fig. 5). Note that the lip in this case does lack all purple coloration. Color forms such as these are not unique in the genus and have been described in H. spicata (forma lutea, wilderi), and H. grandiflora (forma luteoalba) (Brown, 2008).

Fig.1 Hexalectris nitida forma albescens M. White & P.M. Brown


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Hexalectris nitida L.O. Williams forma albescens M. White & P.M. Brown forma nov. TYPE: U.S.A.: Texas, Dallas County. Cedar Hill, Cedar Ridge Preserve off Cedar Break Trail, in oak-juniper woods July 20, 2008. M. White s.n. Fig. 1-3,6. (Holotype: BRIT) Forma plantae et floribus leuteo albescens conspeciebus diversa. Differing from the type in having yellowish-white flowers and stems.

Fig. 2 Cleistogamous flower typical Fig. 3 Flower teased open showing colored lip of the forma albescens

Fig. 4 Distribution of Hexalectris nitida in Texas; with observations of forma albescens indicated by Fig. 5 Hexalectris warnockii forma flavida green dots indicate widespread, red dots local, populations Adapted from Brown & Folsom, Field Guide to the Wild Orchids of Texas, University Press of Florida, 2007.


269

Fig. 6

Photographs by Matt White

Brown: TWO COLOR FORMS FROM THE CENTRAL APPALACHIANS

270

TWO COLOR FORMS FROM THE CENTRAL APPALACHIANS

Paul Martin Brown Color forms in many of our North American native orchid genera are not that unusual and in the genus Corallorhiza color variation is almost the norm. In the recently described Corallorhiza bentleyi J.V. Freudenstein (1999), itself a rarity, a few plants have been observed that are lacking in anythocyans and have pure yellow stems and flowers. These distinctively colored plants are often more colorful than the more typical brownish colored plants that accompany them. Corallorhiza bentleyi J.V. Freudenstein forma flavida P.M. Brown forma nov. TYPE: U.S.A.: West Virginia, Pochahontas County, roadside near junction of routes 250, 28, and Rd. 54 just north of Rd. 54, east side of road by pullout; plants yellow, lip pale yellow, unspotted; 8 plants scattered in woodland. August 9, 2007. P.M. Brown 70809. Fig. 1. (Holotype: WVA) Forma plantae et floribus flavida conspeciebus diversa. Differing from the type in having yellow flowers and stems. Listera smallii Wiegand (1899) is an occasional orchid primarily of the southern Appalachians ranging from disjunct sites in northern New Jersey through the Alleghanies of central Pennsylvania and then becoming more frequent in the mountains of West Virginia south to northern Georgia. When the species was first published (as L. reniformis, a name already in use for an Asian species of Listera) the color of the type specimen was clearly stated as green.

Perennial, fleshy, deep green. Stem erect, 1-3 dm. tall, slender glabrous or nearly so below, densely glandular-pubescent above, simple ; leaves 2, opposite, about the middle of the stem, reniform or ovate-reniform, 1-3 cm. in diameter, apiculate or short acuminate, glabrous above, more or less pubescent beneath, cordate or subcordate, sessile ; racemes 2-10 cm, long ; flowers greenish ; bracts lanceolate or ovate-lanceolate, 3-5 mm. long, acute… (Small, 1897).

Although the richly cinnamon-colored flowers often identified with this species may be more frequent, and often appear in illustrations, they have never be formally treated as a color form. Listera smallii Wiegand forma cinnamomea P.M. Brown forma nov. TYPE: U.S.A.: Pennsylvania, Centre County. ca. 10 mi SSE of State College, Rothrock State Forest, along Bear Meadows Rd., 31 July 1993, leaf litter under Rhododendron. J. Scott Shriver 127 with Albert Shriver, Clete Smith. Fig. 2. (Holotype: CM) Forma floribus cinnamomea conspeciebus diversa. Differing from the type in having cinnamon-colored flowers.

LITERATURE CITED: Freudenstein, J.V. 1999. A New Species of Corallorhiza (Orchidaceae) from West Virginia, U.S.A. Novon 9(4): 511-13. Small, J.K. 1897. Studies in the Botany of the Southeastern United States-XI. Bulletin of the Torrey Botanical Club 24(7): 334. Wiegand, K.M. 1899. A revision of the genus Listera. Bulletin of the Torrey Botanical Club, 26(4): 157-71. Special thanks to Bonnie Isaac, Collection Manager, Carnegie Museum Herbarium (CM), Donna Ford-Werntz, Collection Manager, West Virginia University Herbarium (WVA).

Brown: TWO COLOR FORMS FROM THE CENTRAL APPALACHIANS

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Fig. 1 Corallorhiza bentleyi forma flavida P.M. Brown Pochahontas Co., West Virginia August 10, 2007

Fig. 2 Listera smallii forma cinnamomea P.M. Brown

Centre Co., Pennsylvania 21 July 1996

Carlsward & Whitten: THE CORRECT GENUS FOR THE JINGLE BELL ORCHID, HARRISELLA PORRECTA

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THE CORRECT GENUS FOR THE JINGLE BELL ORCHID, HARRISELLA

PORRECTA

Barbara Carlsward & Mark Whitten

Note: Mark Whitten recently brought to my attention two of his recent publications that validate the inclusion of the familiar Florida jingle-bell orchid, Harrisella porrecta, in the genus Dendrophylax. The following is taken from these two papers: Molecular phylogenetics of neotropical leafless Angraecinae (Orchidaceae): reevaluation of generic concepts. Barbara S. Carlsward, W. Mark Whitten, and Norris H. Williams International Journal of Plant Science 164(1):43–51. 2003 and Ghosts of the Caribbean. Mark Whitten and Barbara S. Carlsward. Orchids 75(10): 742-49. 2006. PMB

All species of Dendrophylax, Harrisella, and Polyradicion are leafless, but Campylocentrum includes

both leafy and leafless species. Flower size varies dramatically among and within genera. Dendrophylax funalis (Sw.) Benth. ex Rolfe, Dendrophylax fawcettii Rolfe, Dendrophylax sallei (Rchb.f.) Benth. ex Rolfe, and Polyradicion lindenii (Lindl.) Garay produce large white, nocturnally fragrant flowers with long, nectariferous spurs (ca. 15 cm in D. fawcettii), whereas other species of Dendrophylax produce much smaller, greenish flowers. Harrisella porrecta (Rchb.f.) Fawc. & Rendle has tiny, greenish tan flowers with a short (ca. 1 mm), bilobed spur; the inflorescence is few flowered and relatively lax. In contrast, most species of Campylocentrum produce short, congested inflorescence axes bearing 10–20 small, white flowers with relatively short spurs. Many of the Old World Angraecinae have large white, spurred flowers that exhibit a hawkmoth pollination syndrome.

Because vegetative characters are reduced or greatly altered in these leafless orchids, the generic concepts of Neotropical Angraecinae have been based largely on gross floral and pollinarium morphology. Most species were originally placed in the genus Aeranthus Lindl. by early workers and were later separated from the Paleotropical taxa into segregate genera. The most recent comprehensive taxonomic treatment of Neotropical Angraecinae is that of Nir (2000) in his examination of Antillean Orchidaceae. In this work, Nir (2000) transferred Polyradicion and Campylocentrum constanzense Garay into Dendrophylax and transferred Harrisella into Campylocentrum, leaving two Neotropical genera distinguished by flower resupination and fruit size.

To date, four species of Harrisella have been described: H. porrecta (Fig. 1,2), Harrisella filiformis (Sw.) Cogn., Harrisella Monteverdi (Rchb.f.) Cogn., and Harrisella uniflora Dietrich. Ackerman (1995) examined the types of these taxa and concluded that H. filiformis and H. monteverdi were synonymous with the leafless Campylocentrum filiforme (Sw.) Cogn. ex Kuntze. Citing unpublished studies of Cuban Harrisella by Jorge Ferro Díaz, Ackerman (1995) also regarded H. uniflora as a synonym of H. porrecta. Conversely, Nir (2000) regarded H. uniflora as a synonym of C. filiforme. In both cases, Harrisella was reduced to a single species, H. porrecta.

Specimens were obtained from cultivated material, herbarium specimens, or wild-collected plants. Samples of Polyradicion lindenii, Campylocentrum pachyrrhizum (Rchb.f.) Rolfe, and Harrisella porrecta from Fakahatchee Strand State Preserve State Park, Florida, and H. porrecta from Grand Cayman are unvouchered; we were only allowed to collect root tips because of the rarity of these species at these localities. Protocols for extraction, amplification, and DNA sequencing from fresh and silica gel–dried material are given in Whitten et al. (2000).


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The results of this work required the transfer of Harrisella porrecta to the genus Dendrophylax.

Dendrophylax porrectus (Rchb.f.) Carlsward & Whitten Basionym. Aeranthus porrectus Rchb.f., Flora 48:279, 1865. Synonyms. Campylocentrum porrectum (Rchb.f.) Rolfe, The Orchid Review 11:247, 1903; Harrisella porrecta (Rchb.f.) Fawc. & Rendle, Journal of Botany 47:266, 1909; Harrisella amesiana Cogn., Symbolae Antillanae 6:687, 1910; Harrisella uniflora H. Dietr., Die Orchidee 33:18–19, 1982. Distribution. El Salvador, Florida, Greater Antilles, Mexico.

Fig.1 Flowers of D. porrectus, showing the short, Fig. 2 D. porrectus (in fruit) from south Florida, growing on saccate, bilobed spur at the base. Taxodium twigs; the roots are intertwined with spun silken tubes of web spinner insects (Embyoptera).

The molecular data also show that the large-flowered D. lindenii (syn. Polyradicion lindenii) and the diminutive D. porrectus (syn. Harrisella porrecta) are embedded within Dendrophylax, and merely represent extremes in flower size and adaptations to different sizes of pollinators (Fig. 3). Most species of Dendrophylax are endemic to the Caribbean island of Hispaniola (Haiti and the Dominican Republic) and bear small, relatively inconspicuous greenish to white flowers. Most are rarely collected by botanists and are poorly known. Botanical exploration of the mountains of the Dominican Republic and Cuba is likely to reveal undescribed species. Dendrophylax porrectus is the most wide-ranging species and can be locally abundant on slender twigs of a wide range of host trees, ranging from the humid swamps of south Florida to seasonally dry


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forests of Yucatan to dry scrub forests in the Dominican Republic. Although the floral morphology of D. porrectus appears relatively uniform across its range, the plants vary geographically in size, root thickness, and habitat preference. DNA data reveal substantial sequence divergence among populations, and we suspect that there may be cryptic species within D. porrectus that have gone unrecognized by botanists because both floral and vegetative morphology are so reduced.

Fig. 3 One of 162 shortest evolutionary trees resulting from a heuristic parsimony search of the combined data matrix for three gene regions (ITS, trnL-F, and matK).

LITERATURE CITED: Ackerman, J.D. 1995. An Orchid Flora of Puerto Rico and the Virgin Islands. New York Botanical Garden, New York. Nir, M.A. 2000. Orchidaceae Antillanae. DAG Media, New York. Whitten, M.W., N.H. Williams, and M.W. Chase. 2000. Subtribal and generic relationships of Maxillarieae (Orchidaceae) with emphasis on Stanhopeinae: combined molecular evidence. American Journal of Botany 87:1842–57. Barbara Carlsward, PhD., Department of Biological Sciences, Eastern Illinois University, 600 Lincoln Avenue, Charleston, IL 61920 email: [email protected] Mark Whitten, PhD., Senior Biological Scientist, 385 Dickinson Hall, Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL 32611-7800 email: [email protected]; Photographs by Mark Whitten Note: For readers who want a simplified explanation of cladistics (how these trees are made and how to read them), see Carlsward: Using the trees to see the forest. The Orchid Review. 111(no. 1251): 166-167, 174, & 178) 2003..

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Lamont & Stalter: ORCHIDS OF ATLANTIC COAST BARRIER ISLANDS FROM NORTH CAROLINA TO NEW YORK

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ORCHIDS OF ATLANTIC COAST BARRIER ISLANDS FROM NORTH CAROLINA TO NEW YORK

Eric E. Lamont & Richard Stalter

Reprinted with permission from Journal of the Torrey Botanical Society 134: 540-551. 2007.

Orchid populations of the Middle Atlantic States are generally not noted as occurring on Atlantic coast barrier islands. Of the 51 orchid species of the Middle Atlantic States discussed by Gupton and Swope (1986) only one, Spiranthes vernalis, is reported as occurring on "sandy beaches". Correll (1950) noted that barrier islands provide suitable habitat for several species of Spiranthes, including S. cernua on "beach sand dunes", S. gracilis on "beaches along the coast", S. praecox on the upper edges of "coastal salt marshes", and S. vernalis on "sandy beaches and dune areas" and "coastal salt marshes"; however, Correll (1950) reported no other orchid species from barrier island habitats. Luer (1975), (1981), Keenan (1998), Brown (2004), and Fowler (2005) included no specific reference to barrier island habitats when discussing habitat preferences of orchid species of the Middle Atlantic States. Duncan and Duncan (1987) included Corallorhiza wisteriana, Cypripedium acaule, and Epidendrum conopseum in their discussion of Seaside Plants of the Gulf and Atlantic Coasts but did not specifically mention barrier islands. Flora of North America (FNA 2002) also made no specific reference to orchid occurrences on Atlantic coast barrier islands but did note that Spiranthes cernua, S. lacera var. gracilis, and S. vernalis occurred on "dunes" and in "dune hollows". Radford et al. (1968) and Gleason and Cronquist (1991) made no reference to barrier island habitats when discussing orchid species within their respective manuals.

During the past 25 years we have been conducting floristic inventories of Atlantic coast barrier islands from North Carolina to New York, and during the course of these studies we have observed and documented the occurrence of 17 orchid species. We have accumulated extensive data on specific localities of extant orchid populations, fluctuations in population size from year to year, flowering dates, and habitat preferences. Although not the focus of this paper, we also have noted potential negative impacts on some populations. In addition to our field studies, we have conducted literature and herbarium searches of historical orchid populations and we have compiled data on orchid populations recently reported by other field botanists but not observed by us.

This paper presents the results of our 25 year study of the orchid flora of Atlantic coast barrier islands from North Carolina to New York. No previous publication has presented a comprehensive study of the current status of orchids occurring on these barrier islands.

MATERIALS AND METHODS Floristic inventories of Atlantic coast barrier islands from North Carolina to New York were

conducted from 1982 to present. Initially, the purpose of these inventories was to document with voucher collections all vascular plant species on major barrier islands and publish the results in botanical journals. To date, six studies have been published, one is in preparation for publication, and another is in plant collection and identification stages.

We initiated our field work in 1982 with a study of the vascular flora of Fire Island, New York (Stalter et al. 1986). In 1985, we extended our survey of New York barrier islands by trying to


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relocate historical orchid populations on Coney Island, Jones Island, Long Beach Island, Rockaway Beach, and Tiana Beach (Lamont et al. 1988, Lamont 1996, 2000). In 1986, we began field studies of selected barrier islands along the Virginia coast, including Assateague Island (Stalter and Lamont 1990), Fisherman Island (Stalter and Lamont 2000a), and False Cape (Stalter et al. 1990, Lamont and Stalter in prep.). Floristic studies of 19 other Virginia barrier and marsh islands (McCaffrey and Dueser 1990, Klotz 1986, Clovis 1968, and Harvill 1965) were consulted and reports of orchid populations have been included in Appendix 1. In 1989, we initiated a 10 year floristic study of the Outer Banks of North Carolina, including Bodie Island, Bogue Banks, Core Banks, Hatteras Island, Ocracoke Island, Pea Island, Portsmouth Island, and Shackleford Banks (Stalter and Lamont 1997, 1999). Our field studies of New Jersey barrier islands have been restricted to Little Beach Island (Stalter 1994) and Sandy Hook (Stalter and Lamont 2000b); however, historical reports of orchid populations in the published literature (Stone 1910, Small and Martin 1958, Martin 1959) and recent reports from New Jersey field botanists have been included in Appendix 1. In 2006, we initiated floristic studies of Delaware barrier islands, including Fenwick Island and the islands comprising Delaware Seashore State Park. We have not conducted any field studies of Maryland's barrier islands because they have been extensively investigated by Hill (1986), Higgins et al. (1971), Reed (1964), Tatnall (1946), and current local and state botanists; however, reports of orchid populations from these sources have been incorporated into Appendix 1.

Because most orchid species are rare to uncommon on Atlantic coast barrier islands, we collected voucher specimens only from large populations usually consisting of 100 or more individuals, although sometimes we collected vouchers from populations of approximately 25 individuals. From smaller populations we either collected one or two individual flowers from an inflorescence or documented the occurrence with photographs. Voucher specimens have been deposited at National Park Service herbaria at each study site, with some duplicates of Spiranthes deposited at the New York State Museum (NYS) in Albany.

During the course of our barrier island studies, we have been informed of additional orchid populations observed and reported by other botanists, but not verified by us. Some of these reports were based upon recent observations and collections by local and/or state field botanists; other reports were historical, based upon publications and collections from the late 1800s to 1980. We have attempted to verify all vouchered reports by examining the original collections and we have cautiously accepted some of the unvouchered reports. In both cases, we have clearly distinguished in Appendix 1 our own personal observations and collections from those of others. Some reports of barrier island orchids published by others have been based upon misidentifications and other published reports are in our opinion questionable; we have included and briefly discussed these incorrect and questionable published reports in Appendix 2.

Appendix 1 presents a summary of the orchid species observed by us on mid-Atlantic coast barrier islands. Species have been arranged alphabetically by genus, followed by common name(s). Location data are presented alphabetically by state, county, name of barrier island, and owner/manager of each site. Habitat data, population size, and blooming period are based upon our own personal observations. Each species entry concludes with miscellaneous comments including but not limited to reports of orchid populations from barrier islands not surveyed by us, pertinent voucher collections examined by us, extirpated populations, and threats to specific populations.


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Appendix 2 presents a summary of the orchid species not observed by us but reported by others from mid-Atlantic coast barrier islands. Species have been arranged alphabetically by genus followed by common name. Location data follows the same sequence as in Appendix 1. The source of each report is given followed by our comments on the status of the species at the reported locality.

Atlantic coast barrier islands change continuously in shape and location, a feature that serves to distinguish these landforms from most others (Ehrenfeld 1990). The majority of study sites included in our survey are currently "island" landforms, but they may have been connected to a mainland in the past; conversely, other study sites are currently contiguous with a mainland, but may have been islands in the past.

Throughout this paper, nomenclature follows the Orchidaceae treatment in Flora of North America (FNA 2002), herbarium abbreviations follow Holmgren et al. (1990), and rarity status of species follows Weakley (2007) and Young (2007). For a detailed description of each study site, see the original sources cited above in the Methods section.

RESULTS AND DISCUSSION

Seventeen orchid species have been documented by us from Atlantic coast barrier islands from North Carolina to New York (Appendix 1). Spiranthes vernalis is the most common species, found at seven localities; Spiranthes cernua was found at five localities; Cypripedium acaule and Listera australis at four; Platanthera cristata, Pogonia ophioglossoides, and Spiranthes odorata at three; Malaxis spicata, Spiranthes praecox, and Tipularia discolor at two; and Calopogon tuberosus, Corallorhiza wisteriana, Epipactis helleborine, Goodyera pubescens, Habenaria repens, Spiranthes lacera var. gracilis, and S. laciniata were each at one locality.

In addition to our own personal observations, Spiranthes vernalis has been reported from 13 other barrier island localities not verified by us; Spiranthes cernua from eight other localities; Calopogon tuberosus and Pogonia ophioglossoides from five other localities; Cypripedium acaule, Platanthera cristata, and Spiranthes lacera var. gracilis from two other localities; and Corallorhiza wisteriana, Goodyera pubescens, Listera australis, and Spiranthes praecox from one other locality (Appendix 1). We have not encountered any additional reports of Epipactis helleborine, Habenaria repens, Malaxis spicata, Spiranthes laciniata, S. odorata, and Tipularia discolor from mid-Atlantic coast barrier islands.

Orchid species occurring as large populations include Platanthera cristata, Spiranthes cernua, S. praecox, S. vernalis, and Tipularia discolor. One population of Platanthera cristata occurring in a moist maritime woodland dominated by Pinus taeda on Assateague Island, Virginia, consists of thousands of individuals; this population is dominated by individuals with pale yellow flowers and may represent forma straminea (Brown et al. 1995). Spiranthes cernua, S. praecox, and S. vernalis are each represented by thousands of widely scattered colonies and individuals in moist to wet sandy soils along roadsides and adjacent swales especially on the Outer Banks of North Carolina. Spiranthes praecox becomes rare on barrier islands north of Chesapeake Bay, whereas S. cernua and S. vernalis remain relatively common. More than a thousand individuals of Tipularia discolor occur in a mature, maritime deciduous forest dominated by Fagus grandifolia and Quercus spp. at Kitty Hawk Woods on


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Bodie Island, North Carolina; in a small study plot of approximately 0.5 ha., we counted 434 individual plants in March 2000.

Four orchid species occur in moderately large populations but have limited distributions. Fifty to more than 100 individuals of Calopogon tuberosus, Cypripedium acaule, and Pogonia ophioglossoides consistently flower at a few barrier island localities in North Carolina and Virginia. Although Epipactis helleborine is a relatively recent introduction to barrier and coastal islands in and near New York Harbor, more than 100 individuals were counted at Sandy Hook, New Jersey.

Orchid species that are rare on mid-Atlantic coast barrier islands include Corallorhiza wisteriana, Goodyera pubescens, Habenaria repens, Listera australis, Malaxis spicata, Spiranthes lacera var. gracilis, S. laciniata, and S. odorata. Each of these species is represented by only a few populations and/or a few individuals.

Several regions of high orchid diversity can be identified on barrier islands from North Carolina to New York. The region between and including Nags Head Woods and Kitty Hawk Woods on Bodie Island, North Carolina, provides suitable habitat for a relatively large number of orchid species, including Cypripedium acaule, Goodyera pubescens, Habenaria repens, Listera australis, Platanthera cristata, Pogonia ophioglossoides, Spiranthes odorata, S. praecox, and Tipularia discolor. False Cape, Virginia, is another orchid "hot spot", providing suitable habitat for Calopogon tuberosus, Cypripedium acaule, Goodyera pubescens, Listera australis, Platanthera cristata, Pogonia ophioglossoides, Spiranthes odorata, S. praecox, and S. vernalis. Assateague Island, Virginia, supports populations of Cypripedium acaule, Listera australis, Platanthera cristata, Spiranthes cernua, and S. vernalis. These orchid hot spots exist because of the high diversity of habitats present at these localities. Few barrier islands north of Assateague Island provide the diversity of habitats required to support large numbers of orchid species.

The most abundant habitat available to orchid species on mid-Atlantic coast barrier islands includes open uplands and wetlands with sandy soils dominated by grasses, sedges, rushes, and forbs. Dry maritime dunes, freshwater interdunal swales, shallow freshwater marshes, and roadsides provide suitable habitat for populations of Calopogon tuberosus, Cypripedium acaule, Habenaria repens, Pogonia ophioglossoides, Spiranthes cernua, S. lacera var. gracilis, S. laciniata, S. praecox, and S. vernalis. Several related yet distinguishable maritime forests cover the more stable portions of barrier islands (Bellis 1995) and provide habitat for populations of Corallorhiza wisteriana, Cypripedium acaule, Goodyera pubescens, Listera australis, Malaxis spicata, Platanthera cristata, and Tipularia discolor.

Three orchid species have been reliably reported by others from mid-Atlantic coast barrier islands, but have not been observed by us in the field (Appendix 2). In 1976, Malaxis unifolia was collected from Nags Head Woods on Bodie Island, North Carolina. We examined the herbarium collection and searched Nags Head Woods in vain for this diminutive orchid (Stalter and Lamont 1997); in our opinion, it is probable that M. unifolia still persists at this locality because suitable habitat is abundant. Stone (1910) reported Liparis loeselii and Platanthera lacera from Long Beach Island, New Jersey. We do not question the validity of these reports, although we have not seen any voucher specimens. In our opinion, it is likely that these populations have been extirpated due to habitat destruction.


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Four orchid species have been reported by others from mid-Atlantic coast barrier islands, based upon misidentifications and misinterpretation of primary sources (Appendix 2). Calopogon pallidus has been reported from False Cape, Virginia, by Knepper et al. (1991), Wright (1991), and Virginia Natural Heritage Program (fide Loomis, pers. comm.), based upon the misidentification of a herbarium collection of Calopogon tuberosus. Krings (2002) reported Platanthera ciliaris as a "new addition to the Outer Banks flora", based upon the misidentification of a herbarium collection of Platanthera cristata. Wright (1991) reported Liparis liliifolia from False Cape, Virginia, but in our opinion this unvouchered report is based upon a misinterpretation of a primary source. Lewis (1917) reported Spiranthes ovalis from Shackleford Banks, North Carolina, but in our opinion this unvouchered report is unlikely and is probably based upon a misidentification of another species of Spiranthes.

In conclusion, during the course of our investigations we have observed an overall decline in the number of orchid populations on barrier islands of the Middle Atlantic States. This decline is largely the result of habitat destruction by humans for development projects. The barrier islands of New Jersey have been most severely impacted resulting in the extirpation of many orchid populations, whereas the barrier islands of Virginia and North Carolina have been less disturbed by human activities and in some regions they remain relatively pristine (e.g., False Cape, Virginia). The federal government's management of ‗national seashores‘ has been the single most significant factor in preserving the ecological integrity of barrier islands of the Middle Atlantic States, and these protected regions often function as refugia for many orchid species.

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Wright, J.B. 1991. A catalog of the vascular flora of the Back Bay watershed, n 257-264. In Marshall, H. G. and M. D. Norman [eds.], Proc. Back Bay Ecol. Symposium. Virginia Beach, Va.

Young, S.M. 2007. New York Rare Plant Status Lists. New York Natural Heritage Program, Albany, N.Y.

Eric E. Lamont, Honorary Research Associate, Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458 Richard Stalter, Department of Biological Sciences, St. John's University, Jamaica, NY 11439 We sincerely appreciate and express thanks to the many individuals acknowledged throughout this paper for assistance in the field, arranging for herbarium loans, checking herbarium label data, and sharing localities of orchid populations not observed by us in the field. Author for correspondence: E-mail: [email protected] APPENDIX 1. Annotated checklist of orchids observed by Lamont and Stalter on Atlantic coast barrier islands from North Carolina to New York Calopogon tuberosus (L.) B.S.P. Common Grass-pink LOCATION: Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, Back Bay National Wildlife Refuge. HABITAT: Wet, sandy, open depressions dominated by sedges, rushes, and forbs. POPULATION SIZE: Approx. 50 to 60 flowering individuals. BLOOMING PERIOD: May to June, sometimes later. COMMENTS: Stone (1910) reported three populations of C. tuberosus from Long Beach Island, Ocean Co., NJ, in the vicinity of North Beach Haven, Spray Beach, and Surf City; we are unaware of any recent efforts to relocate these occurrences. Lamont (1996) cited an herbarium collection [3 Jul 1918, Taylor, BKL] of this species from Tiana Beach, Suffolk Co., NY; extensive attempts by Lamont et al. (1988) to relocate this population were unsuccessful. In 1934 and 1935, Fernald et al. collected C tuberosus from three sites just north of Back Bay National Wildlife Refuge, VA: south of Rudee Inlet [Fernald & Long 3874, HUH], south of Dam Neck, [Fernald, Griscom & Long 4614, HUH], and in swales behind the dunes at Sandbridge [Fernald, Griscom & Long 4615, HUH]. Goldman (pers. comm.) recently collected C. tuberosus from "the False Cape/Back Bay area" (see additional discussion under C. pallidus in Appendix 2). Listed as rare ("Watch List") in VA (Weakley 2007). Corallorhiza wisteriana Conrad Spring Coralroot, Wister's Coralroot LOCATION: North Carolina, Carteret Co., Shackleford Banks, Cape Lookout National Seashore. HABITAT: Low, moist, shady, maritime woodland.

POPULATION SIZE: Few individuals; 11 flowering individuals observed in 1994, none in 1997. BLOOMING PERIOD: April to May. COMMENTS: Apparently, Corallorhiza wisteriana was first reported from Shackleford Banks by Shufun Au (Au 1974). Au also collected this species in 1968 from Harkers Island, Carteret Co., NC (voucher at DUKE) and J. H. Carter collected it in 1972 from Wrightsville Beach, New Hanover Co., NC (voucher at WNC). Luer (1975) considered C. wisteriana to be local and sporadic throughout its range, "being found abundantly one year and scarcely at all the next". Listed as rare ("Watch List") in NC (Weakley 2007). Cypripedium acaule Ait. Pink Lady's-slipper, Pink Moccasin-flower LOCATIONS: New York, Suffolk Co., Fire Island, Fire Island National Seashore. North Carolina, Dare Co., Bodie Island, Nags Head Woods Ecological Preserve. Virginia, Accomack Co., Assateague Island, Assateague Island National Wildlife Refuge; City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. HABITAT: In NC and VA, in dry sands on old stable dunes in maritime forests dominated by Pinus taeda; rarely, as at False Cape State Park, in dry sands under Quercus virginiana. In NY, in dry sands of an open swale between the primary and secondary dunes. POPULATION SIZE: Usually between 30 to 50 widely scattered individuals at Bodie Island and False Cape, approx. 10 individuals at Assateague Island, and 3 individuals at Fire Island. BLOOMING PERIOD: Late April and May in NC & VA, May and June in NY. COMMENTS: McAvoy (pers. comm.) reported C. acaule in 2000 from "sandy pine woods" on barrier


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islands in Sussex Co., DE. Small and Martin (1958) reported C. acaule from Island Beach State Park, Ocean Co., NJ, but Anderson and Kelly (pers. comm.) were unable to relocate this population in 1997. The population at Fire Island, NY, has not been observed since the early 1990s and is considered extirpated, probably the result of selective browsing by white-tailed deer (Odocoileus virginianus). Epipactis helleborine (L.) Crantz Broad-leaved Helleborine LOCATION: New Jersey, Monmouth Co., Sandy Hook, Gateway National Recreation Area. HABITAT: Along disturbed edges of sandy paths in successional woodlands dominated by Prunus serotina. POPULATION SIZE: More than 100 individuals and spreading. BLOOMING PERIOD: Late June to early August. COMMENTS: Also occurs at nearby Ellis Island in New York Harbor (Stalter and Scotto 1999). Although several authors have described E. helleborine as an aggressive weed (Brackley 1985, Soper and Murray 1985, Lamont 1994, Chapman 1997, Brown 2003), it has yet to exhibit such aggressiveness on Atlantic coast barrier islands. Homoya (1993) reported E. helleborine from "sandy, acidic dune forests bordering Lake Michigan" (also see Swink 1966). Introduced from Eurasia (Luer 1975). Goodyera pubescens (Willd.) R. Br. Downy Rattlesnake-plantain, Rattlesnake Orchid LOCATION: Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. HABITAT: Dry sands on old stable dunes in maritime pine-oak forest. POPULATION SIZE: 5 widely scattered individuals observed in 1990. BLOOMING PERIOD: Early July to mid-August. COMMENTS: Reported by Brown (1957) as "rare under beech trees in Nags Head Woods [Bodie Island, Dare Co., NC], identification based upon foliage only", and later collected from the same locality by Gifford [6 Aug 1976, Gifford, CAHA 655]. Extensive attempts by Stalter and Lamont (1997) to relocate this population were unsuccessful. Habenaria repens Nutt. Water-spider Orchid, Floating Orchid LOCATION: North Carolina, Dare Co., Bodie Island, south end of Jockey's Ridge State Park. HABITAT: Wet, sandy, open depression dominated by sedges, rushes, and forbes, including Pogonia ophioglossoides. POPULATION SIZE: Varies from year to year, apparently in response to water level fluctuations;

approx. 15 flowering individuals observed in the early to mid-1990s. BLOOMING PERIOD: May to June, sometimes later. COMMENTS: Near northern limit of its range in Dare Co., NC (FNA 2002). Listed as rare ("Watch List") in NC (Weakley 2007). Listera australis Lindl. Southern Twayblade LOCATIONS: North Carolina, Dare Co., Bodie Island, Kitty Hawk Woods and Nags Head Woods Ecological Preserve. Virginia, Accomack Co., Assateague Island, Assateague Island National Wildlife Refuge); City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. HABITAT: In dense shade and rich humus of low moist maritime forests. At the NC sites the maritime forest is dominated by deciduous species including Quercus spp., Carya glabra and Liquidambar styraciflua; at the VA sites the maritime forest is dominated by Pinus taeda and Quercus spp. POPULATION SIZE: All four populations are relatively small, ranging from 6 to approx. 24 individuals. Several scattered colonies occur within Nags Head Woods. BLOOMING PERIOD: Late April to May. COMMENTS: Gifford collected a voucher specimen [June 1976, Gifford, CAHA] of L. australis from Buxton Woods, Hatteras Island (Cape Hatteras National Seashore), Dare Co., NC; extensive attempts by Stalter and Lamont (1997) to relocate this population were unsuccessful. The small size and inconspicuous nature of this orchid makes it exceedingly difficult to find. Listed as rare ("Watch List") in NC (Weakley 2007). Malaxis spicata Swartz. Florida Adder's-mouth LOCATIONS: North Carolina, Carteret Co., Bogue Banks, Theodore Roosevelt State Natural Area; Dare Co., Hatteras Island, Buxton Woods, Cape Hatteras National Seashore. HABITAT: Low, moist (very wet some years) maritime swamp forest, with Acer rubrum, Quercus laurifolia, Pinus taeda, Saururus cernuus, Pilea fontana, and Boehmeria cylindrica. POPULATION SIZE: Large population at Buxton Woods, some years approximately 100 individuals; usually less than 10 individuals at the Bogue Banks locality. BLOOMING PERIOD: July to mid-August. COMMENTS: Difficult to find due to the inconspicuous greenish brown flowers and apparent preference for inaccessible habitats. Listed as rare ("Active List") in NC (Weakley 2007).


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Platanthera cristata (Michx.) Lindl. Crested Fringed Orchid, Orange-crested Orchid LOCATIONS: North Carolina, Dare Co., Bodie Island, Nags Head Woods Ecological Preserve. Virginia, Accomack Co., Assateague Island, Assateague Island National Wildlife Refuge; City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park near the VA/NC border. HABITAT: Open areas (50 to 70% canopy cover) of sandy, moist maritime woodlands dominated by Pinus taeda with scattered Liquidambar styraciflua and Quercus laurifolia, and Chasmanthium laxum as a dominant groundcover. POPULATION SIZE: One population at Nags Head Woods consisting of approx. 20 to 50+ individuals; two populations at Assateague Island, the northern one consisting of 1000s of individuals, the southern usually with 50 to 100 individuals; one population at False Cape consisting of approx. 50 individuals. BLOOMING PERIOD: July to August. COMMENTS: Reported by Hill (1986) from Assateague Island, MD; Lea (pers. comm.) relocated this population in 1997 and reported approx. 30-40 plants, but in 2000 no plants appeared. In 1934, a voucher specimen of P. cristata was collected from Brier Island, Cape May Co., NJ (5 Aug 1934, Tullner, PH); we are unaware of any recent efforts to relocate this occurrence. The color of mature flowers is usually a deep orange (Luer 1975, FNA 2002), but the large population on Assateague Island, VA, is dominated by individuals with pale yellow flowers. Pogonia ophioglossoides (L.) Ker Gawler Rose Pogonia, Snake's-mouth, Adder's-mouth LOCATIONS: North Carolina, Dare Co., Bodie Island, south end of Jockey's Ridge State Park.Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. New York, Suffolk Co., Fire Island, Smoky Hollow Bog at Fire Island Pines. HABITAT: In NC and VA, occurring in wet, sandy, open depressions with Sphagnum spp., sedges, rushes, and forbes (including Habenaria repens at the NC site); in NY, occurring in a coastal plain poor fen (Edinger et al. 2002) with sedges, rushes, and forbs, bordered by a shrubland community dominated by Clethra alnifolia, Rhododendron viscosum, Vaccinium corymbosum, and Pinus rigida. POPULATION SIZE: 100s of individuals at the NC site, approx. 30 to 50 individuals at the VA site, and approx. 12 individuals at the NY site. BLOOMING PERIOD: May to June in NC and VA; June to early July in NY. COMMENTS: In 1998, Lea (pers. comm.) reported a small population of P. ophioglossoides from Assateague Island, MD, not previously reported by Hill (1986).

Stone (1910) and Small and Martin (1958) reported P. ophioglossoides from Island Beach State Park, Ocean Co., NJ, but Anderson and Kelly (pers. comm.) were unable to relocate this population in 1997. Stone (1910) also reported P. ophioglossoides from Long Beach Island, Ocean Co., NJ, in the vicinity of North Beach Haven and Spray Beach; we are unaware of any recent efforts to relocate these occurrences. Lamont (1996) cited an herbarium collection [13 Jul 1918, Taylor, BKL] of this species from Tiana Beach, Suffolk Co., NY; extensive attempts by Lamont et al. (1988) to relocate this population were unsuccessful. During the mid- 1990s, increased groundwater use by seasonal residents at Fire Island Pines, NY, contributed to a decrease in the local water table resulting in the succession of Smoky Hollow Bog into a shrubland community; in 2000, no P. ophioglossoides was found at the site. Listed as rare ("Watch List") in VA (Weakley 2007). Spiranthes cernua (L.) Rich. Nodding Ladies'-tresses LOCATIONS: Delaware, Sussex Co., Fenwick Island, Fenwick Island State Park. New York, Suffolk Co., Fire Island, Sailor's Haven and Watch Hill/ Bayberry Dunes at Fire Island National Seashore. North Carolina, Dare Co., Hatteras Island, roadside near Buxton. Virginia, Accomack Co., Assateague Island, Assateague Island National Wildlife Refuge. HABITAT: In DE, NC, and VA occurring along moist, sandy roadsides and adjacent swales in full sun; in NY occurring in moist, open, interdunal swales often with Vaccinium macrocarpon and in a freshwater marsh. POPULATION SIZE: Usually 100s, some years 1000s, of widely scattered individuals at each location. BLOOMING PERIOD: Late August to October, sometimes later in NC. COMMENTS: Spiranthes cernua becomes more common on barrier islands north of Chesapeake Bay. McAvoy (pers. com.) reported it "on interdunal swales on barrier islands in Sussex Co., DE on Rehoboth and Indian River Bays." Hill (1986) reported several occurrences on Assateague Island, Worcester Co., MD; Lea (pers. comm.) relocated these populations in 1999. Reed (1964) reported S. cernua "on sand dunes" in the vicinity of Ocean City, Fenwick Island, Worcester Co., MD. Small and Martin (1958) reported S. cernua from Island Beach State Park, Ocean Co., NJ, but Anderson and Kelly (pers. comm.) were unable to relocate the population in 1997. Stone (1910) reported three populations of S. cernua from Long Beach Island, Ocean Co., NJ, in the vicinity of Harvey Cedars, Ship Bottom, and Spray Beach; we are unaware of any recent efforts to relocate these occurrences. Stone (1910) also reported S. cernua from Sandy Hook, Monmouth Co., NJ, but Stalter and Lamont (2000b) were unable to relocate the population.


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Spiranthes lacera (Raf.) Raf. var. gracilis (Bigel.) Luer Southern Slender Ladies'-tresses LOCATION: New York, Suffolk Co., Fire Island, Fire Island National Seashore, east of Sailor's Haven. HABITAT: Dry sands in full sun, bayside of interdunal swale. POPULATION SIZE: One population, approx. 20+ individuals. BLOOMING PERIOD: August to September, sometimes later. COMMENTS: Reed (1964) cited an herbarium collection [1 Sept 1932, Redmond, DOV] of S. lacera var. gracilis from Fenwick Island, Worcester Co., MD; we are unaware of any recent efforts to relocate this occurrence. Lamont (1996) cited two herbarium collections [26 Aug 1915, Bicknell, NY; 27 Aug 1916, Pennell, NY] of this species from Long Beach, Nassau Co., NY; extensive attempts by Lamont et al. (1988) to relocate this population were unsuccessful. Correll (1950) reported, "this species is found in . . . beaches along the coast . . . near sea level along the Atlantic and Gulf coasts"; however, we have found this species to be very rare on Atlantic coast barrier islands from NC to NY. Spiranthes laciniata (Small) Ames Lace-lipped Ladies'-tresses LOCATION: North Carolina, Carteret Co., Cape Lookout National Seashore. HABITAT: Shallow freshwater marsh (in standing water at least part of the year) occurring with graminoids and forbs, including a large population of Sagittaria lancifolia var. media (= S. falcata). POPULATION SIZE: One small population, less than 10 flowering individuals. BLOOMING PERIOD: July (we have seen reports of this species also flowering in NC during June and early Aug, but from 1994 to 1997 we observed the population at Cape Lookout National Seashore consistently flowering during July). COMMENTS: The voucher specimen of S. laciniata collected by us (consisting of a few flowers and a leaf) from Cape Lookout National Seashore was determined by Charles Sheviak (NYS). Spiranthes laciniata can be distinguished from S. vernalis, which it superficially resembles, by its capitate trichomes; it typically flowers later than S. vernalis where the two are sympatric (FNA 2002). Listed as rare ("Active List") in NC (Weakley 2007). Spiranthes odorata (Nutt.) Lindi. Fragrant Ladies'-tresses LOCATIONS: North Carolina, Dare Co., Bodie Island, vicinity of Kitty Hawk. Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, Back

Bay National Wildlife Refuge and False Cape State Park. HABITAT: In NC occurring in swampy areas often dominated by Quercus laurifolia and Morella [Myrica] cerifera; in VA along the edges of Back Bay tidal creeks and channels and in tidal marshes. POPULATION SIZE: In NC, scattered colonies each with usually less than 20 individuals; in VA, several dozen individuals concentrated in the region of Major Cove and adjacent Big Ball Island, Horse Island Creek, and Deal Creek. BLOOMING PERIOD: October to early November. COMMENTS: Spiranthes odorata is a southeastern coastal plain endemic (Weakley 2007) and is near the northern limit of its range at False Cape, VA. Listed as rare ("Active List") in VA (Weakley 2007). Spiranthes praecox (Walt.) S. Wats. Giant Ladies'-tresses, Grass-leaved Ladies'-tresses LOCATIONS: North Carolina, Dare and Currituck Cos., Cape Hatteras National Seashore north to the VA line. Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, Back Bay National Wildlife Refuge. HABITAT: Disturbed, wet sands in full sun, thrives along roadsides where periodic mowing discourages competition. POPULATION SIZE: 1000s of individuals widely distributed throughout the Outer Banks of NC; dozens of individuals at False Cape, VA. BLOOMING PERIOD: June and July. COMMENTS: Spiranthes praecox is a southeastern coastal plain endemic (Weakley 2007) and is rare on Atlantic coast barrier islands north of Chesapeake Bay. McCaffrey and Dueser (1990) did not find S. praecox on any of Virginia's 16 barrier islands just north of Chesapeake Bay, nor did Stalter and Lamont (1990) find it on Assateague Island, VA. Hill (1986) found a single population on Assateague Island, MD, but Lea (pers. comm.) could not relocate it in 1996 and 1997. Stalter and Lamont (pers. obs.) found no populations of S. praecox on Delaware barrier islands in 2006, and Stone (1910) did not report it from any New Jersey barrier islands. Spiranthes vernalis Engelm. & Gray Spring Ladies'-tresses LOCATIONS: Delaware, Sussex Co., Delaware Seashore State Park; Fenwick Island, Fenwick Island State Park. New York, Suffolk Co., Fire Island, Fire Island National Seashore. North Carolina, Carteret Co., Bogue Banks, vicinity of Fort Macon State Park; Currituck Co., Bodie Island, vicinity of Corolla Light. Virginia, Accomack Co., Assateague Island, Assateague Island National Wildlife Refuge; City of Virginia Beach (formerly Princess Anne Co.), False Cape, Back Bay


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National Wildlife Refuge; Northampton Co., Fisherman Island, Fisherman Island National Wildlife Refuge. HABITAT: Disturbed, wet sands in full sun, thrives along roadsides where periodic mowing discourages competition; also in freshwater marshes and wet, interdunal swales with graminoids and forbs, often associated with Vaccinium macrocarpon. POPULATION SIZE: Each locality usually comprised of widespread, small populations or widely scattered individuals; total number of individuals at each site varies from 100s or 1000s to approx. 10. BLOOMING PERIOD: May (in NC and se VA) to July (early August in NY). COMMENTS: Spiranthes vernalis is the most common orchid on Atlantic coast barrier islands from NC to NY. Hill (1986) reported it as "widespread" on Assateague Island, MD, and Lea (pers. comm.) relocated it there in 1997 and reported it as "fairly common". Reed (1964) cited an herbarium collection [11 Jul 1931, Redmond, DOV] of S. vernalis from "behind sand dunes, Ocean City", Fenwick Island, Worcester Co., MD; we are unaware of any recent efforts to relocate this occurrence. Stone (1910) reported S. vernalis as being "rather frequent along the coast strip" of NJ and listed specific occurrences from Atlantic Co. (Absecon Island, vicinity of Atlantic City and Longport), Cape May Co. (Five Mile Beach Island, vicinity of Wildwood, and Ocean City barrier island), and Ocean Co. (Long Beach Island, vicinity of Beach Haven Terrace and Spray Beach). In 2000, Snyder (pers. comm.) reported extant NJ populations of this orchid "in wet to damp swales behind sand dunes" from Atlantic Co. (Brigantine Island), Cape May Co. ("several locations on barrier islands"), and Ocean Co. (Long Beach Island, vicinity of Holgate). Lamont (1996, 2007) cited two herbarium collections [4 Aug 1911, McCallum, BKL; 27 Jul 1922, Bicknell, NY] of this species from Coney Island, Kings Co., NY and Long Beach, Nassau Co., NY, respectively; extensive attempts by Lamont et al. (1988) to relocate these

populations were unsuccessful. Klotz (1986) and McAvoy (1996, pers. comm.) reported S. vernalis from Accomack Co., VA, in disturbed sands on Wallops Island and moist sandy swales on Parramore Island, respectively. McCaffrey and Dueser (1990) reported it from Hog Island and Smith Island, Northampton Co., VA, based upon field work conducted in 1975; in 1996, McAvoy (pers. comm.) relocated the Hog Island population in moist sandy soil at the island's north end. Listed as rare ("Active List") in NY (Young 2007). Tipularia discolor (Pursh) Nutt. Crane-fly Orchid LOCATIONS: North Carolina, Dare Co., Bodie Island, Kitty Hawk Woods. Virginia, Northampton Co., Fisherman Island, Fisherman Island National Wildlife Refuge. HABITAT: In NC occurring in the thin layer of humus of a mature, maritime deciduous forest dominated by Quercus falcata, Q. nigra, and Fagus grandifolia with an understory of Ilex opaca and Cornus florida, on low, gently undulating, old, stable dunes; in VA occurring in thin humus of a mesic maritime woodland community dominated by Prunus serotina with a few scattered individuals of Ilex opaca. POPULATION SIZE: 1000s of individuals in Kitty Hawk Woods (in a small study plot of approx. 0.5 ha., we counted 434 individual plants in March 2000); one small population of 7 individuals on Fisherman Island. BLOOMING PERIOD: Mid-July to August. COMMENTS: Tipularia discolor readily colonizes woodlands in the early stages of development or regrowth (Homoya 1993); such conditions are characteristic of the locality on Fisherman Island, VA. It is notoriously difficult to locate flowering individuals of T. discolor, "so well does their leafless and inconspicuous flowering stalks blend into the sticks and leaves and filtered sunbeams" (Luer 1975); the best time of year to locate this species is during midwinter when the leaves are conspicuous.

APPENDIX 2. Annotated checklist of orchids not observed by Lamont and Stalter but reported by others from Atlantic coast barrier islands from North Carolina to New York Calopogon pallidus Chapman Pale Grass-pink LOCATION: Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, Back Bay National Wildlife Refuge. SOURCE OF REPORT: Knepper et al. (1991); Wright (1991); Virginia Natural Heritage Program (fide Loomis, pers. comm.). These reports were based upon a voucher specimen, originally identified as Calopogon

pallidus and deposited at Old Dominion University Herbarium, with the following label data: "Back Bay, Black Gut, Sphagnum swamp, Sandbridge bog, 10 June 1957, V. Bagley s.n., ODU 23508". [Note: although C. pallidus was reported from Back Bay National Wildlife Refuge by Virginia Natural Heritage Program, Black Gut is located just north of the refuge.] COMMENTS: The above cited Bagley collection at ODU was examined by Douglas H. Goldman (HUH)


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on 22 Jan 2004 and was reassigned to Calopogon tuberosus. Goldman (pers. comm.) commented: "To date [Aug 2006], I've been through 148 herbaria surveying Calopogon and I've never seen any C. pallidus specimens from the False Cape/Back Bay area. I've been in that area before and the only species collected there was C. tuberosus, which is supposedly locally common in that area. I have a plant list from False Cape State Park from 1/13/91 and it lists C. pallidus, but that is almost surely incorrect (Calopogon, where species are sympatric, seem to be commonly misidentified). The 1992 Atlas of the Virginia Flora does not list this species [C. pallidus] as being from Virginia Beach City Co. or Chesapeake City Co., only Isle of Wright Co. and Suffolk City Co.". Liparis liliifolia (L.) Rich, ex Lindi. Lily-leaved Twayblade LOCATION: Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. SOURCE OF REPORT: An unvouchered report by Wright (1991) gleaned from an earlier report by Fernald (1935). COMMENTS: Fernald (1935) began the fifth paragraph of Midsummer Vascular Plants of Southeastern Virginia by noting, "Farther south, in the damp depressions in the sand between Back Bay and the dunes of False Cape is another area of localized coastal plain plants .. .". The next paragraph begins, "The rich woodlands, as already noted by Mr. Griscom and me, contain many species of the interior most unexpected on the outer edge of the coastal plain, close to the Atlantic. Our browsings brought to light many other plants of rich woods, several of them amazingly remote from their inland centers: Thelypteris hexagonoptera (Michx.) Weath., Panicum Boscii Poir., Liparis liliifolia (L.) Richard ... Yet in the woods near Little Creek it is less than a mile from the sea." At first reading, it may appear that Fernald reported L. liliifolia from False Cape; however, Fernald's voucher collection confirms the locality as "Little Neck, Princess Anne Co., Virginia" [Fernald, Griscom & Long 3869, HUH]. Later, Fernald (1936) identified Little Neck as "projecting into Lynnhaven Bay", on the mainland just west of Cape Henry. We therefore conclude that Wright's (1991) report of L. liliifolia from False Cape State Park is based upon a Fernald et al. collection and report from the nearby mainland. Additionally, we have not seen a voucher specimen of L. liliifolia collected from False Cape State Park, nor does the park provide suitable habitat to support a population of L. liliifolia. Liparis loesellii (L.) Rich. Loesel's Twayblade LOCATIONS: New Jersey, Atlantic Co., Absecon Island; Ocean Co., Long Beach Island. New York,

Nassau Co., Long Beach; Queens Co., Rockaway Beach. SOURCE OF REPORTS: Stone (1910) reported one occurrence of L. loesellii from the southern tip of Absecon Island in the vicinity of Longport, and two occurrences from Long Beach Island in the vicinity of Beach Haven Terrace and Surf City. Lamont (1995, 1996) cited two herbarium collections [26 Aug 1900, Bicknell, NY; 27 Jun 1896, Mulford, BKL] of this species from Long Beach, Nassau Co., and Rockaway Beach, Queens Co., respectively. COMMENTS: We are unaware of any recent efforts to relocate the three NJ occurrences reported by Stone (1910). Extensive attempts by Lamont et al. (1988) to relocate the two NY occurrences were unsuccessful. In our opinion, it is likely that these populations have been extirpated due to habitat destruction. Malaxis unifolia Michx. Green Adder's-mouth LOCATIONS: North Carolina, Dare Co., Bodie Island, Nags Head Woods Ecological Preserve. Virginia, City of Virginia Beach (formerly Princess Anne Co.), False Cape, False Cape State Park. SOURCE OF REPORTS: A voucher specimen collected from Nags Head Woods [12 May 1976, Gifford, CAHA], and an unvouchered report by Wright (1991) from False Cape State Park gleaned from an earlier report by Fernald (1935). COMMENTS: In our opinion, it is probable that M. unifolia still persists at Nags Head Woods, but extensive efforts to relocate it have been unsuccessful (Stalter and Lamont 1997). Fernald (1935) collected M. unifolia "in rich woods, Back Bay, Princess Anne Co., Virginia" [Fernald, Griscom & Long 4622, HUH], but we cannot determine with certainty if the collection was from False Cape. The "Back Bay" citation by Fernald (1935) may refer to the town of Back Bay located on the adjacent mainland just west of Drum Point (see additional discussion under comments for Liparis liliifolia). Fernald (1935) did specifically cite "False Cape" as the locality for many of his collections (e.g., Pogonia ophioglossoides). Platanthera ciliaris (L.) Lindi. Yellow Fringed Orchid LOCATION: North Carolina, Dare Co., Bodie Island, Nags Head Woods Ecological Preserve. SOURCE OF REPORT: Krings (2002) reported Platanthera ciliaris as a "new addition to the Outer Banks flora", based upon a voucher specimen [6 Aug 1976, Gifford s.n., CAHA 657] deposited in the herbarium of the National Park Service Cape Hatteras National Seashore Unit, Manteo, NC. The specimen was originally identified by Gifford as Habenaria ciliaris, and


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in 2001 Krings annotated the collection and updated the name to Platanthera ciliaris. COMMENTS: We examined the above cited Gifford collection and reassigned it to Platanthera cristata. Morphological differences in flower structure (e.g., spur length and rostellum shape) clearly distinguish P. cristata from P. ciliaris (FNA 2002). A population of P. cristata was located at Nags Head Woods by Stalter and Lamont (1997). It is our opinion that P. ciliaris should be excluded from the orchid flora of Atlantic coast barrier islands from NC to NY. Platanthera lacera (Michx.) G. Don in R. Sweet Ragged Fringed Orchid LOCATION: New Jersey, Ocean Co., Long Beach Island. SOURCE OF REPORT: Stone (1910) reported two occurrences of P. lacera from Long Beach Island in the vicinity of Beach Haven Terrace and Holgate. COMMENTS: We are unaware of any recent efforts to relocate these two occurrences. In our opinion it is likely that these populations have been extirpated due to habitat destruction. Spiranthes ovalis Lindl. Oval Ladies'-tresses LOCATION: North Carolina, Carteret Co., Cape Lookout National Seashore, Shackleford Banks. SOURCE OF REPORT: An unvouchered report by Lewis (1917). COMMENTS: Lewis' (1917) report of Spiranthes ovalis from Shackleford Banks contains some peculiarities. Spiranthes ovalis is first mentioned on page 25 of his

publication, following a list of "ten species [from Shackleford Banks] which are stated by Small not to occur north of South Carolina". The ten species are listed alphabetically and annotated with range data. Immediately following the list is an indented paragraph that begins with: "Spiranthes ovalis has also not been reported from this State." Why wasn't S. ovalis included in the original list of new state records? Then on page 28, in the section of the publication that lists the plants of Shackleford Banks, Lewis includes a single taxon under the Orchidaceae: "Spiranthes ovalis Lindl. (?); (Gyrostachys parviflora (Chapm.) Small (?))." Lewis' question marks give the impression that he was uncertain about his identification. On page 11, Lewis states: "Exact determination of the plants found is of the highest importance in such an account as this. This has been made possible by the kindness of Dr. John K. Small, of the New York Botanical Garden, to whom specimens of practically all the plants listed were sent." We could not find Lewis' Spiranthes collection at The New York Botanical Garden (NY), nor has it been deposited at the UNC- Chapel Hill herbarium (fide McCormick, NCU). Extensive efforts by Stalter and Lamont (1999) to relocate S. ovalis on Shackleford Banks were unsuccessful. Correll (1937), Radford et al. (1964), and Weakley (2007) all reported a restricted range for S. ovalis in NC, limited to the piedmont physiographic province. In our opinion, Lewis' (1917) report is probably based upon a misidentification of another species of Spiranthes. According to Sheviak and Brown (FNA 2002), only S. ovalis var. erostellata Catling occurs in NC and adjacent states.

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THE FUTURE…?

The Slow Empiricist There are more and more tools being discovered to help the orchid enthusiast learn more

about native orchids. There are people working with DNA to unlock the lineage and discover family connections. Some people are studying the mycorrhizal configurations and influence in the life of the different species. People also study habitat, pollinators, and climate to learn more about our native orchids. These endeavors should bode well for survival and propagation of native species even as civilization encroaches on habitat and pollution and all of the man made ills threaten some species with extinction. I have many concerns about the carelessness of human activity that impacts not only our native orchids but all aspects of the natural world. Our penchant for wanting the newest, biggest, or best innovation of the ‗must-have‘ generation is frightening. Small homes in older desirable neighborhoods are bought up, torn down, filling up landfills and new McMansions are installed in their place requiring more material to be taken out of nature‘s supply to fulfill this ‗need‘. Even less affluent people have the remodeling bug; they must have granite countertops or marble bathrooms. Tear out and replace! The sad thing is in 10 or 15 years these ‗must-haves‘ will be old hat and dated in need of replacement. I could give lots of other examples but I think the intelligent reader already knows the excesses people go to in order to look better, richer, or have more than their neighbors. Therefore, I will concentrate on what this mindset might do in the native orchid world. If people become successful in the quest to identify the mycorrhizal connections orchid species rely upon to grow, will that enable us to grow these orchids in our gardens and greenhouses? Will this lead to the hybridization of the natives to produce better plants? ….ones that are hardier or have a longer blooming period. Or will some enterprising entrepreneur work to improve the look of the flower? I can imagine Spiranthes with grossly exaggerated lips becoming the preferred Spiranthes than its normal looking ancestors. I can see highway departments beautifying the roadsides with these grotesque plants which have been bred to be hardier than their native ancestors and their eventual domination in the habitat to the detriment of the original species. An example is highway beautification projects where the hardier, larger Western ox-eye daisy has been introduced—on eastern roadsides. It is overwhelming the less-showy Eastern ox-eye daisy and may soon dominate and crowd out the native variety. I have seen what orchid societies in this country have done with the exotic orchids they are so enamored with. The Phalaenopsis orchids have lost their starry shapes to fulfill some arbiter‘s


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vision that proclaims they should be as round as a saucer. Again, I won‘t belabor this treatise with the fashion of color that breeders work for or the garishness that collectors look for to the detriment of the less opulent native only to say it is a sad state of affairs. Hopefully the future for native North American orchids will assure the enthusiast that the integrity of the species won‘t be compromised to fashion or to other conceived human needs. I hope we will be wise enough to use the new tools at hand to ensure the continued survival of the orchids as nature intended not as man wants. Your Slow Empiricist


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BOOK REVIEWS ORCHIDS OF RUSSIA AND ADJACENT COUNTRIES (WITHIN THE BORDERS OF THE FORMER

USSR). Vakhrameeva, M.G., I.V. Tatarenko, T.I. Varlygina, G.K. Torosyan, and M.N. Zagulski. 2008. 97 color distribution maps. 233 photographs and watercolor plates. ix + 690 pp. 8vo. Cloth. English. ISBN 978-3-906166-61-2. A.R.G. Gantner Verlag, Ruggell, Liechtenstein. €148.00. Distributed by Koeltz Scientific Books, Koenigstein, Germany https://www.koeltz.com. Order inquiries to Orchid Books (CAD 244.95; ca. USD 228.95), http://www.orchidsbooks.com/ or directly to Koeltz.

The Orchids of Russia….. is a massive work that covers so much of Eastern Europe and northern Asia and treats nearly all of the temperate terrestrial orchids to be found in that region. It is the first complete modern scientific treatment of the orchid flora of the former USSR. The authors studied orchids for more than 30 years, in different climatic zones—polar regions of the Cola Peninsula, temperate zones in Central Russia, Central Ukraine, Baltic Sea coast territories, Mediterranean regions in the Crimea and Caucasus, continental climate in Siberia and Central Asia, and monsoonic climate in the Far East of Russia. The first several chapters consist of detailed essays by the various authors entitled History and Contemporary Tendencies in Orchid Studies, Systematic Review, Species Distribution, Ecology and Phytocoenology, Morphology, Ontogenesis, Rhythm of Seasonal Development and Reproduction, Mycorrhiza, and Species Protection. The bulk of the book is devoted to the species descriptions and they are very complete and easy to understand. Throughout the book extensive use of references is made resulting in more than 70 pages in the bibliography. A brief glossary contains not only the technical terms but several Anglicized Russian terms that help in understanding the text. I now know that a Zapovendik is a highly protected area under government control that prohibits economic activities or a ‗nature preserve‘.

A work such as this is written over many years and judging from the references contains data through 2004. Several genera and species also found in North America are included, many of these quite rare and found especially in Alaska. It is helpful to have more information on these species. They include Calypso bulbosa, Cypripedium guttatum, C. yatabeanum, Dactylorhiza aristata, Goodyera repens, Liparis loeselii, Listera cordata, Lysiella oligantha (Platanthera obtusata subsp. oligantha), Malaxis monophyllos, Platanthera convallariifolia, P. dilatata, P. chorisiana, and P. tipuloides. Several of our non-native North American species are native in this region and include treatments of Epipactis atrorubens, E. helleborine, E. palustris, Listera ovata, and Zeuxine strateumatica. One of the most interesting treatments is in the use of the genera Lysiella as mentioned above and Tulotis. The latter would include the eastern North American Platanthera flava.

Many scientific works originally written in a language other than English often suffer greatly in translation but not so with this book. The English is excellent and the proofreading exceptional. Over 200 color photos of the species are complemented by numerous excellent watercolors and supplemented by 18 line drawings and 93 distribution maps. It is here that the book has its only significant fault. No key map is provided for either the geography or showing the several ecological regions so often mentioned in the book. For those not intimately familiar with the many countries of the former USSR and their locations this can be very confusing. If only one page were devoted to a simple outline map showing the borders of these countries it would make the book much more usable. Fortunately such a map is easily accessed on the Internet and I found an excellent one at www.theodora.com. The names of a few of the countries are slightly different from those used in the book, but easily understood. The map can be printed out and inserted within the book for

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reference. I highly recommend Orchids of Russia for all orchid professionals and enthusiasts with an interest in the native orchids of this region. PMB

THE MARIE SELBY BOTANICAL GARDENS ILLUSTRATED DICTIONARY OF ORCHID GENERA Peggy Alrich & Wesley Higgins; Bruce Hansen (Editor); Robert L. Dressler (Editor); Tom Sheehan (Editor); John Atwood (Editor); Peter H. Raven (Foreword) Cornell University Press. A Comstock Book $49.95, cloth, 2008, xxv + 482 pages, 8.5 x 11 in., full-color illustrations throughout, ISBN: 978-0-8014-4737-2. Published in association with Selby Botanical Gardens Press

It is not often that such a compendium of orchid information is assembled in so attractive a user-friendly format. The The Marie Selby Botanical Gardens Illustrated Dictionary of Orchid Genera presents its information in such a way that it becomes accessible to both the novice orchid enthusiasts and dedicated professional alike. The authors and editors should be congratulated for their work. If paid by the hour they would all be millionaires from this work alone!

From the Profile of the Orchid Family by David Benzing, which could easily stand alone as one of the best examples of its kind, through the extensive Glossary that concludes the volume all of the information is presented in such a way that is interesting, captivating, and informative. In several places the book states specifically that this is an ongoing project and taxonomic opinions are subject to revision. The Dictionary presents those of a suite of authors and editors ultimately reflecting the opinion of the Orchid Identification Center of the Marie Selby Botanical Gardens through the end of 2007. BUT that is what synonyms are for.

Because of the arrangement of the entries it is easy to find virtually any genus that has been used for an orchid for well over 400 years! I have been pleasantly surprised with any number of obscure genera that host a number of synonyms. The generic treatments are such that they are color coded, and the codes are repeated on bottom of the pages throughout the generic treatments, which makes is much easier to quickly determine if the generic name is in current usage, a synonym, invalid, or even a fossil record. Individual flowers are shown for many of the genera. The taxonomic treatment is interspersed with wonderful images from classic orchid works, many from early herbals. Following the generic descriptions are a full list of taxonomists who have made generic descriptions and revisions, a detailed list of publication abbreviations cited, and a concise presentation of the International Rules of Botanical Nomenclature governing the valid publication of names. The latter is especially appreciated and will be useful for many enthusiast who may not have knowledge of, or access to, this information. An extensive list of the illustrations by page number is given and the credits for the graphics grouped together on a single page. Although it would have been unwieldy to credit each flower graphic, it would have been appreciated to credit some of the original art work for the book. Artists often suffer this way. The final page is a request for feedback in the way of errors, omissions, and new generic information.

A work such as massive as this is not without is problems, and although most are minor it is embarrassing to see that the first photograph in the book, Fig. 5, is labeled Harrisella porrecta, when the genus Harrisella is treated within the genus Dendrophylax in the text as per recent work by Carslward et al. (2004).

Our North American natives always seem to suffer in larger works, especially when information is gleaned from older publications. Although I in no way profess to have a universal knowledge of orchid genera I am familiar with the North America genera and species north of Mexico and have found a number of curious statements or errors. Most of these in no way detract from the usefulness of the work. Amerorchis (p. 18) is stated as currently included in the genus Platanthera but I can find no current publications that do so. The range cited is also incorrect, as

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Amerorchis is documented for New York, Vermont, and Maine. Arethusa (p. 33) also occurs in South Carolina. Blephariglottis (p. 48) is illustrated with what is identified as Blephariglottis albiflora (syn. Platanthera blephariglottis) but is not that species and is a pale colored flower of P. grandiflora. The description of the species within the genus Blephariglottis does not include all of the purple-flowered species. Dichromanthus (p. 123) extends into Arizona. Eburophyton, (p. 141) as Cephelanthera austiniae, is well-documented from Salt Spring Island, in southwestern British Columbia, Canada and in the treatment of Cephalanthera is noted from British Columbia. Fimbriella (p. 161) is cited as including a single species, F. (Platanthera) lacera, but the IPNI includes six species, five of those in the original paper describing the genus. The illustration is identified as F. lacera, but appears to be a curious artist‘s composite of several species and not clearly identifiable as to any one, especially not to F. lacera. The range give for F. lacera omits Texas. Gymnadeniopsis (p. 181) is cited as having an uppermost lip in the three species and that is incorrect. Only two of the three species have the lip in this position. Habenaria (p. 183) curiously omits the southeastern United States in the range, where this genus is particularly abundant. I was particularly interested to see the genus Hammarbya (p. 185) used as most North American authors still include it in Malaxis although in Eurasia the latter is in general usage. The range given omits central Canada – Manitoba and western Ontario. Isotria, (p. 204) extends into northern Florida and eastern Texas. The genus Listera (p. 225), although considered by the authors now in the genus Neottia, extends south to central Florida. Microthelys (p. 246) has been found in New Mexico, a new record for the Untied States, in 2004. Orchis as a valid genus (p. 278) is stated as occurring in North America. Unless one includes the synonyms using this genus, Orchis as we know it today does not occur in North America. The closest, historically, would be the treatment of Orchis spectabilis as Galearis and Orchis rotundifolia as Amerorchis. Paliris (p. 287) is cited as a synonym for Liparis loeselii and the range given omits North America where it is perhaps more abundant than in Eurasia. Pelexia (p. 294) is well documented for Florida. Peramium (p. 295), as a synonym for Goodyera, extends into Georgia, South Carolina, northern Florida, and west to eastern Texas. Piperia (p. 305) is cited as occurring in Russia, although the new Orchids of Russia (2008) does not include this genus (or species). The range given omits New Mexico, South Dakota, eastern Canada and Michigan. Both Pleurothallis (p. 310) and Stelis (p. 372) are listed for southern Florida when it can only be one or the other, but not both. Restrepiella (p. 334) and Tetramicra are (p. 387) both listed for Florida, but it is a well-known and documented fact that they were erroneously included. The inclusion of these two genera is old information and should not have happened. Virtually all, excepting the record of Microthelys, of the distributional information and errors could have been easily fact-checked and corrected by consulting the Orchidaceae in Flora of North America North of Mexico, vol. 26.

Readers should not let these minor points deter them and I can highly recommend this book to all orchid enthusiasts. It is a volume that will be read for pure pleasure and repeatedly used for reference and research. Given the costs of publishing today it is a real buy for $49.95 (and perhaps less if you shop around). PMB

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NEW TAXA AND COMBINATIONS PUBLISHED IN THE NORTH AMERICAN NATIVE ORCHID JOURNAL VOLUME 14, 2008

Volume 14 (1): 1- 67 issued January 15, 2008. Volume 14 (2): 68-184 issued April 3, 2008. Volume 14 (3): 185-235 issued July 15, 2008. Volume 14 (4): 236-292 issued October 15, 2008. Calopogon ×floridensis P.M. Brown nothosp. nov. p. 176 Calopogon ×fowleri P.M. Brown nothosp. nov. p. 177 Calopogon ×goethensis P.M. Brown nothosp. nov. p. 178 Calopogon ×obscurus P.M. Brown nothosp. nov. p. 176-77 Calopogon ×simulans P.M. Brown nothosp. nov. p. 177 Calopogon ×vulgaris P.M. Brown nothosp. nov. p. 177 Corallorhiza bentleyi J.V. Freudenstein forma flavida P.M. Brown f. nov. p. 270 Galearis spectabilis (L.) Raf. forma lilacina (Ames) P.M. Brown stat. & comb. nov. p. 138 Hexalectris nitida L.O. Wiliams forma albescens P.M. Brown f. nov. p. 267 Liparis smallii Wiegand forma cinnamomea P.M. Brown f. nov. p. 270 Platanthera shriveri P.M. Brown sp. nov. p. 239 Platanthera ×enigma P.M. Brown nothosp. nov. p. 255 Sacoila paludicola (Luer) P.M. Brown stat. nov. p. 194 Sacoila paludicola Luer forma aurea (P.M. Brown) P.M. Brown comb. nov. p. 198 Spiranthes stellata P.M. Brown, L.A. Dueck & K.M. Cameron sp.nov. p. 4

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NEW FROM

CORNELL UNIVERSITY PRESS & SELBY BOTANICAL GARDENS PRESS A Comstock Book $49.95, cloth, 2008, xxv + 482 pages, 8.5 x 11 in., full-color illustrations throughout, ISBN: 978-0-8014-4737-2

The Marie Selby Botanical Gardens Illustrated Dictionary of Orchid Genera is the most comprehensive and extensively illustrated account of orchid genera to date. Its concise entries provide details of nomenclature,

classification, original publication, etymology, and geographic range, along with a brief description and color images of representative flowers. The dictionary describes not only all of the 850 orchid genera that are recognized today but also those genera known only from fossil records, published before Linnaeus, validly published (but not accepted), and invalidly published according to the standards of the International Code of Botanical Nomenclature, as well as those that have variant names or spellings. In addition to the alphabetic entries, this dictionary includes an introduction to orchid biology, a glossary, a list of taxonomists credited with publishing new orchid genera, key references and bibliographical abbreviation list, and the governing nomenclature rules. The Marie Selby Botanical Gardens Illustrated Dictionary of Orchid Genera also features a Foreword by Peter H. Raven and an Introduction on the biology of orchids by David Benzing that describes the August 2007 discovery of the world's oldest unequivically orchidaceous fossil. Order your copy from: Cornell University Press http://www.cornellpress.cornell.edu/cup_detail.taf?ti_id=5285 Marie Selby Botanical Gardens http://www.selby.org/index.php?submenu=Research&src=gendocs&ref= Orchid%20Genera&category=Research Orchid Books http://www.orchidsbooks.com/book.asp?id=1833 or your local bookseller

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NEW! from

Gantner Verlag and

Koeltz Scientific

Books Orchids of Russia and Adjacent Countries (within the borders of the former USSR). 2008. 97 color distribution maps. 233 photographs and water color plates. ix + 690 pp. 8vo. Cloth. English. ISBN 978-3-906166-61-2. €148.00.

Distributed by Koeltz Scientific Books, Koenigstein, Germany https://www.koeltz.com. Order

inquiries to Orchid Books http://www.orchidsbooks.com/ or directly to Koeltz.

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or directly from the author at [email protected]

december 2008 north american native orchid journal

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