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NORTH AMERICAN NATIVE ORCHID JOURNAL
______________________________________ Volume 4 March Number 1 1998
a quarterly devoted to the orchids of North America published by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE
* * * * * * *
* * * * * * *
IN THIS ISSUE: POLLINATION BIOLOGY IN SOME MEMBERS OF THE YELLOW FRINGED ORCHID COMPLEX CYPRIPEDIUM ARIETINUM BROWN AND CYPRIPEDIUM PLECTROCHILUM FRANCHET
CHECKLIST OF THE ORCHIDS OF NORTH AMERICA And more…………
NORTH AMERICAN NATIVE ORCHID JOURNAL
(ISSN 1084-7332) published quarterly in
March June September December by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE, Inc.
a group dedicated to the conservation and promotion of our native orchids
Editor: Paul Martin Brown
Assistant Editor: Nathaniel E. Conard Editorial Consultants:
Philip E. Keenan Stan Folsom
Production Assistant: Nancy A. Webb
The Journal welcomes articles, of any length, of both a scientific and general interest nature relating to the orchids of North America. Scientific articles should conform to guidelines such as those in Lindleyana or Rhodora. General interest articles and notes may be more informal. Authors may include line drawings, and/or black and white photographs. Color inserts may be arranged. Please send all inquiries or material for publication to the Editor at PO Box 772121, Ocala, FL 34477-2121 (mid June - August: PO Box 759, Acton, ME 04001-0759). 1999 Membership in the North American Native Orchid Alliance, which includes a subscription to the Journal, is $26 per year for United States addresses, $29US in Canada and $32US other foreign countries. Payment should be sent to Nancy A. Webb, 84 Etna St. Brighton, MA 02135-2830 USA. Claims for lost issues or cancelled memberships should be made within 30 days.
NORTH AMERICAN NATIVE ORCHID JOURNAL
Volume 4 March Number 1 1998
CONTENTS NOTES FROM THE EDITOR
1
POLLINATION BIOLOGY IN SOME MEMBERS OF THE YELLOW
FRINGED ORCHID COMPLEX Part 1. Floral morphology, pollinators
and pollination mechanisms C. P. Argue
3
CYPRIPEDIUM ARIETINUM BROWN AND CYPRIPEDIUM PLECTROCHILUM FRANCHET
H. Perner 31
NEW TAXA P. M. Brown
45
SETTING GOALS
The Slow Empiricist 54
CHECKLIST OF THE ORCHIDS OF NORTH AMERICA
north of Mexico 61
ANNOTATIONS TO THE CHECKLIST P.M. Brown
100
LOOKING FORWARD June 1998
113
ANNOUCEMENT 3rd Annual North American Native Orchid
Conference 114
Unless otherwise credited, all drawings in this issue are by Stan Folsom
Color Plates:
1. p. 41 Cypripedium arietinum 2. p. 42 Cypripedium plectrochilum 3. p. 47 Cypripedium kentuckiense forma pricei; 4. p. 48 Eulophia alta ; Eulophia alta forma pelchatii 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 4, number 1, pages 1- 115; issued March 20, 1998. Copyright 1998 by the North American Native Orchid Alliance, Inc.
Cover: Cypripedium arietinum by Stan Folsom
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POLLINATION BIOLOGY IN SOME MEMBERS OF THE YELLOW
FRINGED ORCHID COMPLEX
Part 1. Floral morphology, pollinators and
pollination mechanisms
Charles P. Argue Recent studies in the yellow fringed orchid complex have resolved some important questions about sexual reproduction in several members of this group. The present paper reviews the results for yellow fringed orchid, Platanthera ciliaris (L.) Lindley and white fringed orchid, P. blephariglottis (Willdenow) Lindley. More limited data are available and summarized for several other species of the complex. Part two, dealing with breeding systems, reproductive success, and selection, will appear in a later issue. Platanthera ciliaris and P. blephariglottis Floral morphology. The plants of Platanthera blephariglottis and P. ciliaris are about 0.10 to 1 meter tall (Smith & Snow, 1976; Folsom, 1984). Their racemes are cylindrical or ovoid in shape, 5 to 15
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cm long and about 5 cm wide; the flowers are bilaterally symmetrical and white in P. blephariglottis or orange in P. ciliaris (Smith & Snow, 1976). The perianth is comprised of broadly oval sepals, 5 to 10 mm long; shorter linear to oblong lateral petals with lacerate distal ends; and an 8 to 16 mm long, tongue shaped, linear to oblong labellum or lip with a fringed margin. The labellum is projected into a downward extending, basal nectar spur which is up to 3 cm long and is often three-quarters filled with nectar (Smith & Snow, 1976; Robertson & Wyatt, 1990a; Cole & Firmage, 1984). Both orchids have an immobile, two celled anther (Luer, 1975). The anther is separated into two half anther sacs located above and to either side of the opening to the spur (Fig. 1a). Each encloses a granular, club-shaped pollinium, which is attached by a narrow stalk to a sticky adhesive pad or viscidium (Smith & Snow, 1976; Robertson & Wyatt, 1990a; Luer, 1975). The pollinia are made up of numerous, small, loosely associated packets of pollen, the massulae, within which individual pollen grains are attached to one another by means of minute elastic threads. The anther locules diverge and the viscidia point upward and outward from the flower (Fig. 1a, b) (Smith & Snow, 1976; Folsom, 1984). One viscidium is located on either side of the opening to the nectar spur. The stigma is directly above the entrance to the spur and
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Figure 1. Floral morphology and pollination in Platanthera ciliaris. Sketch adapted from figures 3 and 8 in Folsom (1984). A. The column in face view. Note position of viscidia. Scale line about 5 mm. B. Lateral view showing long nectar spur and contact between viscidia and eye of butterfly. Scale line about 20 mm. Abbreviations: a = anther, e = eye of butterfly, l = lip of orchid, n = nectar spur, o = nectar spur opening, p = proboscis of butterfly, s = stigma, v = viscidium. See text for explanation.
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beneath and between the half-anther cells (Fig. 1a) (Smith & Snow, 1976; Robertson & Wyatt, 1990a). Pollination mechanisms and pollinators. The pollination mechanism is similar in both species. The insect locates the spur opening and inserts its proboscis. The viscidia are positioned so as to contact and adhere to the compound eyes of the vector (Fig. 1b) (Smith & Snow, 1976; Robertson & Wyatt, 1990a, b), and one or both pollinaria are extracted from the half anther cells as the insect withdraws from the flower (Smith & Snow, 1976). As the stalk dries on one side it rotates down and inward, positioning the pollinium directly to the front of the vector’s head where it may later contact the stigmas of subsequently visited flowers (Luer, 1975; Cole & Firmage, 1984). Pollinaria may remain attached to the eye for several days (Robertson & Wyatt, 1990a, b) and in Platanthera blephariglottis, at least, the pollen can remain viable for upwards of five days (Cole & Firmage, 1984). When the pollinia are brushed against a stigma, individual massulae often separate from one another, and one to several can be deposited on successive stigmas as the pollinator moves from flower to flower (Luer, 1975; Smith & Snow, 1976). In most cases several flowers are explored on the same raceme before the insect moves on (Smith & Snow, 1976). A critical consideration is the relative length of the spur and proboscis. If, for example, the proboscis is distinctly longer than the tube, the insect can
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extract the nectar without contacting the viscidia with its eyes. The spur length in both orchids implicates long-tongued pollinators (Smith and Snow, 1976). Orange to yellow flower color is known to attract a variety of vectors including butterflies, whereas white flowers are often associated with moth pollination (Smith & Snow, 1976; van der Pijl & Dodson, 1966; Faegri & van der Pijl, 1979). However, Cole and Firmage (1984) related white flower color in Platanthera blephariglottis to butterfly pollination, the contrast making the flowers conspicuous against the darker color of the surrounding bog mat and vegetation. These authors also reported the production of only a weak nocturnal as well as diurnal fragrance in P. blephariglottis. This weak diurnal fragrance is a feature not usually associated with moth pollination. However, other floral characters in addition to the white to cream-colored petals and sepals, including an absence of both nectar guides and an ultraviolet pattern, are consistent with pollination by moths (Cole & Firmage, 1984; Faegri & van der Pijl, 1979). Accumulating data now suggest that the association between both orchid species and their pollinators may be complex and include factors dependent upon microclimatic conditions as well as expected geographical differences. Studies are available on P. ciliaris chiefly from Michigan and the Carolinas and for P. blephariglottis from Michigan and Maine.
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Michigan
Based on two years of research conducted at Booth Lake Bog, Berrien County, Michigan, Smith and Snow (1976) concluded that moths were the chief pollinators of Platanthera blephariglottis and butterflies were the chief pollinators of P. ciliaris at this site. According to these authors, butterflies were initially attracted to P. ciliaris by the bright orange color of its flowers; as they approached they were thought to recognize the form of the flower and finally its odor. Nocturnal moths, on the other hand, were said to be first drawn to P. blephariglottis by the flower’s odor and only later, at close range, by flower form and color. In mixed groups of P. ciliaris and P. blephariglottis Smith and Snow (1976) reported that night-flying moths were attracted to the white flowers of the latter but would nevertheless follow an odor trail to isolated groups of P. ciliaris. This differed from diurnal moths which were not seen to visit P. ciliaris (Smith & Snow, 1976). Although contradicted by studies that report little or no floral odor in Platanthera blephariglottis (Cole & Firmage, 1984) and P. ciliaris (e.g., Folsom, 1984), these observations are consistent with van der Pijl and Dodson’s (1966) assertion that at least some diurnal moths (Macroglossa) are more sensitive to color than night-flying species. They are also consistent with a set of observations by Smith and
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Snow (1976) which compared the mean number of flowers of P. ciliaris pollinated in two microhabitats. In the first the racemes were located in open areas, free of concealing vegetation. In the second they were surrounded and partly obscured by other plants. The number of flowers pollinated on each raceme in the first habitat was approximately twice that of the second. Such a result would be expected if a visual stimulus were responsible for attracting the pollinator (Smith & Snow, 1976). For P. blephariglottis approximately the same number of flowers per raceme were pollinated in the two habitats. If the pollinator in this case were attracted by odor, the presence of surrounding vegetation would have had little influence on pollination. Platanthera ciliaris was most frequently pollinated by Papilio troilus L. (spicebush swallowtail) in the Michigan study, although this species did occasionally also visit P. blephariglottis (Smith & Snow, 1976). Its chief larval food, Linderia benzoin (L.) Blume (spicebush), was common in the bog, and the emergence of its second (summer) brood correlated with the peak flowering period for P. ciliaris which occurred one to two weeks later than that for P. blephariglottis (Smith & Snow, 1976). Less frequent pollinators carrying pollinaria of P. ciliaris in Michigan included Papilio glaucus L. (tiger swallowtail), Strymon liparops Boisduval & Leconte (striped hairstreak), and Danaus plexippus L. (monarch). Strymon liparops only rarely had pollinaria attached. Its chief larval food plant, the blueberry
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(Vaccinium), was common in the bog, and its July emergence occurred during anthesis of P. ciliaris (Smith & Snow, 1976). Thus, it might be expected to play a role, but even though its proboscis is of sufficient length to reach the nectar, its head is small and may not always come into contact with the viscidia (Smith & Snow, 1976). Hyles lineata L. (white-lined sphinx), a nocturnal species, was seen to visit the flowers and was considered a probable pollinator by Smith and Snow (1976) although it was not observed carrying pollinaria. The ruby-throated hummingbird, also seen at the flowers, was not considered a pollinator of P. ciliaris because of its long beak and large size (Smith & Snow, 1976).
Smith and Snow’s (1976) data on the pollinators of Platanthera blephariglottis did not reveal a single vector with a predominance comparable to that of the spicebush swallowtail for P. ciliaris. The most frequent insects found carrying pollinaria,
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Platanthera ciliaris orange fringed orchis
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Platanthera blephariglottis white fringed orchis
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each observed three times, were spicebush swallowtails (Papilio troilus) and bumblebees (Bombus sp.) (Smith & Snow, 1976). Nevertheless, these authors considered moths to be the primary pollinators, and those found with attached pollinaria included Darapsa versicolor Harr. (hydrangia sphinx) and Hemaris thysbe L. (hummingbird clearwing). Both were daytime visitors and expected residents of the bog, as the larval foods of these species (Cephalanthus occidentalis L. [buttonbush] for the sphinx moth and Lonicera sp. [honeysuckle] and Viburnum sp. [“cranberry”] for the clearwing) were present in the bog (Smith & Snow, 1976). In addition, a single Danaus plexippus (monarch) was observed with pollinaria attached. As was the case with P. ciliaris, this species was probably no more than an incidental pollinator of P. blephariglottis. It traveled over a wide area, and its larval foods were not found in the bog (Smith & Snow, 1976). Other insects classified by Smith and Snow (1976) as possible to probable pollinators, but not observed with attached pollinaria, included Apis mellifera L. (honeybee) and three night flying moths: Manduca quinquimaculata Haw. (five-spotted hawkmoth), Agrotis sp. (cutworm moth), and Hyles lineata (white-lined sphinx). Apis mellifera has a short proboscis, and its attempts to locate the nectary were described as awkward by Smith and Snow (1976). It was considered an unlikely pollinator. The larval food plants of Manduca quinquimaculata (Lycopersicum and Nicotiana) are not bog plants, and
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it most likely did not visit Platanthera blephariglottis regularly (Smith & Snow, 1976). Similarly, Smith and Snow (1976) considered that Agrostis is probably too small to be effective in transferring pollen. Hyles lineata may be expected as a frequent night visitor since Epilobium (willow herb), one of its larval food plants, occurred in the bogs (Smith & Snow, 1976). As was the case for P. ciliaris, it carried no pollinaria, but Smith and Snow (1976) again believed it to be a likely pollinator. Activity of nocturnal moths may have been inhibited by low night time temperatures in the bog. Night flying moths were inactive when temperatures dropped to 15 C (Smith & Snow, 1976). Only nine nights out of the 28 day flowering period were warm enough for night-flying moth activity (Smith & Snow, 1976). Daytime temperatures were never low enough to interfere with the activity of butterflies, bees or diurnal moths. The percentage of flowers pollinated in Platanthera blephariglottis was inversely related to the size of the raceme. This, according to Smith and Snow (1976), was because each pollinator spent about the same amount of time on large and small racemes. In P. ciliaris, on the other hand, the smaller racemes, those with 10 or fewer flowers, showed a lower percentage of pollination than larger racemes. Smith and Snow (1976) suggested that butterflies, depending on sight to find the orchid, may have overlooked racemes with fewer flowers present. In addition, the mean number of flowers in each
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raceme was higher for P. ciliaris than for P. blephariglottis, a feature which Smith and Snow (1976) also believed may be due to the visual basis of butterfly attraction, resulting in the selection of larger racemes. The percentage of Platanthera ciliaris flowers pollinated (45.5%) was about twice that for P. blephariglottis. According to Smith and Snow (1976), this may have been related to the cool night time temperatures noted above. However, Heath and Adams (1967), in a study of the physiology of the sphinx moth Hyles lineata, reported that this species could, through activity, maintain body temperatures at 34 to 38 C against air temperatures of 10 to 30 C. Based on the number of flowers of Platanthera blephariglottis pollinated on the bottom, middle, and upper thirds of the raceme, Smith and Snow (1976) concluded that pollination rates were more or less constant throughout the blooming period. These authors believed that pollinator numbers were essentially constant throughout, and that the pollinators, chiefly moths attracted by scent, could detect the presence of a few flowers at the bottom of the raceme about as well as a fully blooming raceme. On the other hand, they reported that pollination was 55% greater in the top third as compared to the bottom third of the raceme in P. ciliaris. From this they concluded that pollinator numbers increased in parallel with the number of
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open flowers, and fully blooming racemes were more readily detected by the pollen vectors. Maine More recently, Cole and Firmage (1984), in a three year study at the Colby Marston Preserve, a bog in Kennebec County, Maine, significantly expanded the list of known pollinators for Platanthera blephariglottis. In contrast to the results of the Michigan study, these authors found butterflies to be much more important than moths and the almost exclusive pollen vectors of this species in their study area. Both small and large butterflies were observed as pollinators throughout the flowering period, but some seasonal change was noted with the former being relatively more abundant early and the latter increasing toward the end of the season (Cole & Firmage, 1984). Seventy-nine percent of the visitors and pollinators of P. blephariglottis at this site were butterflies and skippers and 15% were bees. Nearly half of all insects observed on the flowers (46%) were true skippers (Hesperiidae), and nearly a quarter (22%) were of a single species, Epargyreus claurs Crammer (silverspotted skipper). The remainder were primarily whites and sulfurs (Pieridae) and hymenoptera (Apidae). Altogether nine species of lepidoptera and two species of hymenoptera were identified as pollinators (Table 3 in Cole & Firmage, 1984).
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No moths were seen with attached pollinaria during the three year study. Microclimatic differences in the form of cold air drainage sometimes reduced night time temperatures in the Maine bog to below 15 C which may again have inhibited nocturnal moth activity (Cole & Firmage, 1984). However, light-trapping and observation on warm nights also failed to reveal the presence of moths with attached pollinaria (Cole & Firmage, 1984). Two species which Smith and Snow (1976) classified as pollinators, Papilio troilus (spicebush swallowtail) and Danaus plexippus (monarch), were listed as flower visitors by Cole and Firmage (1984). Apis mellifera (honeybee), considered an unlikely but possible pollinator by Smith and Snow (1976), was regarded as a floral visitor by Cole and Firmage (1984). Hemaris and Bombus were thought to be pollinators by both Smith and Snow (1976) and Cole and Firmage (1984). Hemaris was observed visiting the flowers only once in the Maine study, and as already noted, none were found carrying pollinaria (Cole & Firmage, 1984). Cole and Firmage (1984) considered the role of bumblebees in the pollination of Platanthera blephariglottis to be minor (Cole and Firmage, 1984); their tongues are too short for the spur. Bumblebees were implicated in the pollination of Kalmia angustifolia L. (sheep laurel) which stopped flowering before P.
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blephariglottis. Bombus then switched over to P. blephariglottis. In fact, except for this overlap in certain years, the latter provided the only significant source of nectar during its blooming period. Thus, unlike the Michigan site where P. ciliaris was common (Smith & Snow, 1976), butterflies at the Maine site would often have been dependent on P. blephariglottis during its blooming period. Butterflies, however, had little competition for the available nectar, and, in particular, no moth competition except for the infrequently seen diurnal moth Hemaris (Cole & Firmage, 1984). Cole and Firmage (1984) recorded a pattern of pollinator movement on the inflorescence of Platanthera blephariglottis which may differ from that usually observed in Lepidoptera (Wyatt, 1982). The pollinators frequently began their explorations of the inflorescence with the middle to lower flowers and fed toward the top. Usually they visited only a few flowers, feeding on an average of 3.4 (1-19) from the same part of the inflorescence and spending an average of 10 (1.3-39) seconds on each flower (Cole & Firmage, 1984). The average amount of time the pollinator remained on an inflorescence of P. blephariglottis, 34 + 8.2 (2-145) seconds, was less than the mean time required for the stalk of the pollinarium to rotate into position to contact the stigma (about 60 seconds) and effect pollination in that inflorescence (Cole & Firmage, 1984).
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In contrast to Smith and Snow’s (1976) report of constant pollination rates on the bottom, middle, and top thirds of the inflorescence, Cole and Firmage (1984) found significant differences in these rates, with that for the top third being either higher or lower than that for at least one of the other thirds in each of the three years of their study. Thus, some factor or combination of factors such as the number of pollinators or the level of pollinator activity varied throughout the season at the Maine study site. Similarly, Cole and Firmage (1984) were unable to confirm any consistent negative correlation between raceme size and percentage capsule set, except for a weak one during the first year of the study when a relatively large number of racemes were in bloom. Cole and Firmage (1984) also saw little difference in percentage capsule set between orchids growing on open mat or hidden among shrubs or trees. However, they considered that although these plants may have been difficult to locate visually from ground level, most would not have been obscured from potential pollinators flying over them. In any case, butterflies had no difficulty in locating and pollinating orchids surrounded by vegetation (Cole & Firmage, 1984). North and South Carolina In a two year study Robertson and Wyatt (1990a, b) found marked differences in pollination
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ecology between two widely separated populations of Platanthera ciliaris occurring in the coastal plain of South Carolina and the Appalachian mountains of western North Carolina. The coastal plain site was located close to Awendaw in the Francis Marion National Forest, the mountain population at Corveeta Hydrologic Laboratory. Although the species differed, the pollinators of primary importance at both locations were large butterflies (Robertson & Wyatt, 1990a, b). No other insects carried pollinaria and no other insects were observed as consistent visitors (Robertson & Wyatt, 1990a). As in Michigan, the most important pollinator in the mountains of North Carolina during both years of the study was Papilio troilus (spicebush swallowtail) (Robertson & Wyatt, 1990a, b). It was the most frequently observed visitor and usually had pollinaria attached to its eyes; each carried an average of 5.9 to 6.8 over the two years (Robertson & Wyatt, 1990a, b). Battus philenor L. (pipevine swallowtail), a reportedly toxic species (Howe, 1975), is similar to and was sometimes lumped with Papilio troilus, its putative mimic. However, Battis phelinor proved to be a far less frequent pollinator than Papillio troilus (Robertson & Wyatt, 1990a, b). Papilio glaucus (tiger swallowtail) was observed visiting Platanthera ciliaris both years and carried pollinaria (three times) during one year of the study (Robertson & Wyatt, 1990a, b). Phoebis sennae L. (cloudless sulfur) was noted only once. It
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had a single pollinarium attached (Robertson & Wyatt, 1990a, b). On the other hand, the most frequent pollinator on the coastal plain of South Carolina both years was Papilio palamedes Drury. (palamedes swallowtail); over 80% of those examined had pollinaria attached, and each carried an average of about 3.5 (Robertson & Wyatt, 1990a, b). This species is restricted to the southeastern United States and Mexico (Howe, 1975), and its range does not extend to the mountain population of P. ciliaris. Phoebis sennae (cloudless sulfur) was also seen to visit coastal plain populations of Platanthera ciliaris during both years of the study, but only about one-third carried pollinaria, and the average number of pollinaria carried per individual (2.5) was lower than that observed for Papilio palamedes (Robertson & Wyatt, 1990a, b). Within an area where both butterflies were active simultaneously the majority of palamedes swallowtails bore pollinaria whereas only a minority of cloudless sulfurs did (Robertson & Wyatt, 1990a, b). Papilio troilus, the predominant pollinator in the mountains, was occasionally observed on the coastal plain but did not commonly visit Platanthera ciliaris, although one year it was recorded three times, once with a single pollinarium attached (Robertson & Wyatt, 1990a, b). It was often seen on Liatris graminifolia Willd. (blazing star) at a site near the location of Platanthera ciliaris. These flowers were also frequently visited by
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Papilio palamedes and Phoebis sennae (Robertson & Wyatt, 1990a, b). Major pollinators on the coastal plain carried fewer pollinaria than did the pollinators in the mountains, and pollinator activity, evaluated in terms of the rate of deposition and removal of pollinaria, was lower on the coastal plain than in the mountains. Paralleling these observations, percentage fruit set and the total number of fruits per plant were lower on the coastal plain than in the mountains both years of the study. As will be discussed in the second part, the length of the nectar spurs in mountain population of Platanthera ciliaris (23.8 mm) closely approximated the lengths of the probosci in their primary (19.5-24.9 mm) and secondary (19.5-24.9 mm) pollinators. In coastal plain populations, on the other hand, spur length (25.6 mm), although already significantly longer than in mountain populations, was shorter than the probosci of either the primary or secondary pollinators (both ca. 28.7 mm), and selection for longer nectar spurs may be occurring at this site. However, reciprocal transplant studies were inconclusive (Robertson & Wyatt, 1990a), and further research is needed to clearly establish the genetic basis of the differences in floral morphology between mountain and coastal plain populations. Experiments conducted to test for nocturnal pollination found none occurring at either site
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(Robertson & Wyatt, 1990a, b). Xylophanes tersa L. (tersa sphinx moth) carried no pollinaria but stole nectar from the flowers of Platanthera ciliaris on the coastal plain. Seen on only one occasion, this species hovered from plant to plant just after dark and extracted nectar with its long proboscis (32.0 mm long in one individual compared to a spur length of about 24 to 26 mm) without contacting the viscidia (Robertson & Wyatt, 1990a). Sometimes other apparent nectar thieves, possibly carpenter bees, made slits near the bottom of the spurs at both sites and extracted most of the nectar (Robertson & Wyatt, 1990a, b). In the mountains neither sphinx moths nor other nectar thieves with long tongues were observed (Robertson & Wyatt, 1990a). Again, moth activity could have been limited by low temperature, as night time minimums during the flowering season averaged 14.2 C as compared to 20.5 C on the coastal plain (Robertson & Wyatt, 1990a). If visits of sphingid moths at the coastal-plain site are frequent enough, and if they are indeed absent or rare in the mountain population, they could extract sufficient nectar at the former site to effect some difference in the frequency of pollinator visits at the two sites (see part 2) (Robertson & Wyatt, 1990a). Other yellow fringed orchids
A limited amount of information is available on the pollination of at least two other species of
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the yellow fringed orchid complex, Platanthera cristata (Michx.) Lindl. (crested fringed orchid or orange-crested orchid) and P. chapmanii (Small) Luer emend Folsom (Chapman’s fringed orchid). Platanthera cristata is smaller than P. blephariglottis and P. ciliaris and has orange to yellow flowers (Folsom, 1984). Platanthera chapmanii, intermediate in many ways between P. cristata and P. ciliaris and long considered a hybrid of these species, is treated as a distinct species by Folsom (1984). In addition to morphological differences (Folsom, 1984 and below), P. chapmanii and P. cristata are separated from one another and other members of the complex by pollination biology (Folsom, 1984).
In a study conducted on the coastal plain of Florida and Alabama (Folsom, 1984), Platanthera chapmanii, like P. ciliaris, was found to be pollinated by long-tongued butterflies: Papilio troilus, P. palamedes, P. marcellus Cramer, and Phoebis sennae. It differs, however, from the butterfly-pollinated species discussed earlier in the manner of pollinaria attachment (Folsom, 1984). It will be recalled that the length of the spur in P. ciliaris approximately equals the length of the vectors’ probosci, that the anther locules diverge, and that the viscidia point upward and outward from the flower and attach to the eyes of the pollinator (Folsom, 1984). However, the spur in P. chapmanii is only 10 to 14 mm long (Folsom, 1984), and due to a bend in the column, the viscidia converge and face the
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Figure 2. Floral morphology and pollination in Platanthera chapmanii. Sketch adapted from figures 3 and 8 in Folsom (1984). A. The column in face view. Note position of viscidia. Scale line about 5 mm. B. Lateral view of flower with shortness of spur slightly exaggerated. Note positioning of viscidia and contact between viscidia and proboscis of butterfly. Scale line about 10 mm. Abbreviations as in figure 1. See text for explanation.
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Platanthera cristata orange crested orchis
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labellum (Fig. 2a, b). This positioning and the short nectary result in an attachment of the pollinaria to the butterfly’s proboscis rather than to its eyes (Folsom, 1984).
The small flowers of Platanthera cristata have
an even shorter spur, 5 to 8 mm long (Folsom, 1984). The viscidia are closely spaced and face forward. Their alignment and positioning are said to be adapted for attachment of the pollinia to the head of a bee, and it is reportedly pollinated chiefly by the bumblebee, Bombus pennsylvanica Degeer (Folsom, 1984).
To be continued. Literature Cited Cole, R. F. and D. H. Firmage. 1984. The floral ecology of
Platanthera blephariglottis. Amer. Jour. Bot. 71: 700-710. Faegri, K. and L. van der Pijl. 1979. The principals of pollination
ecology. 3rd ed. Pergamon, Oxford. Folsom, J. P. 1984. A reinterpretation of the status and
relationships of taxa of the yellow fringed orchid complex. Orquidea 9: 321-346. Reprinted (1995) in North American Native Orchid Journal 1: 213-238.
Heath, J. E. and P. A. Adams. 1967. Regulation of heat production by large moths. Jour. Expr. Biol. 47: 21-33.
Howe, W. H.. 1975. The butterflies of North America. Doubleday, Garden City, NY.
Luer, C. A.. 1975. The native orchids of the United States and Canada. New York Botanical Garden. Bronx, NY.
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Robertson, L. J. and R. Wyatt. 1990a. Evidence for pollination ecotypes in the yellow fringed orchid, Platanthera ciliaris. Evolution 44: 121-133.
______. 1990b. Reproductive biology of the yellow fringed orchid, Platanthera ciliaris. Amer. Jour. Bot. 77: 388-398.
Smith, G. R. and G. E. Snow. 1976. Pollination ecology of Platanthera (Habenaria) ciliaris and P. blephariglottis (Orchidaceae). Bot. Gaz. 137: 133-140.
van der Pijl, L. and C. H. Dodson. 1966. Orchid flowers: their pollinators and evolution. University of Miami Press, Coral Gables, FL.
Wyatt, R. 1982. Inflorescence architecture: How flower number, arrangement, and phenology affect pollination and fruit-set. Amer. Jour. Bot. 69: 585-594
Charles P. Argue, Ph.D., Department of Plant Biology, University of Minnesota, 220 Biological Sciences Center, 1445 Gortner Avenue, St. Paul, MN 55108-1095. Dr. Argue last wrote for the Journal in December of 1995 on the pollination biology of Arethusa, Pogonia and Calopogon. Figures 1 & 2 drawn by Dr. Argue after Folsom, 1984.
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CYPRIPEDIUM ARIETINUM BROWN
AND CYPRIPEDIUM PLECTROCHILUM
FRANCHET
Holger Perner
We had been traveling for several hours on our way to the northwestern part of New York State, up to the St. Lawrence River in the area of Lake Ontario. Here my friend Dr. Charles ("Chuck") J. Sheviak, curator for botany at the New York State Museum in Albany, wanted to show me the ram's-head lady's-slipper, Cypripedium arietinum, which is rare in the US. After lunch in a small restaurant we went to an area which is called "limestone barrens". Once there we were greeted by remarkably big swarms of black flies and mosquitoes, because usually nobody gets lost there to give them blood. The completely flat area—a former lakebed—showed a uniform substrate of limestone. There were deep gaps between the big flat blocks, some of them small, some of them over 0.5 m wide. A small layer of humus allowed only a few plants to grow, some of it was dropseed, Sporobolus heterolepis (a prairie grass), some of it was denser, and of several species which made up a diverse carpet together with field juniper, Juniperus communis (creeping and erect). There were deciduous
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shrubs and woods dominated by northern white cedar, Thuja occidentalis. The rocks were eroded, concave and created shallow, flat, moist areas.
For a few hours we walked through this area and I
would have been completely lost without Chuck. Occasionally we found a single large yellow lady's-slipper, Cypripedium parviflorum var. pubescens, damaged by frost, and Chuck apologized for bringing me here. Completely an error, for I was excited about the rich flora including such interesting examples as rattlesnake fern, Botrychium virginianum, showy trillium, Trillium grandiflorum, bloodroot, Sanguinaria canadense and much more. Taxonomic History
For a little while I would like to leave the North American wilderness and enter the dusty study rooms of botany and talk about the problem of Cypripedium arietinum and its sister species C. plectrochilum of China.
In 1813 Robert Brown described Cypripedium arietinum of North America in "Hortus Kewensis". Five years later Rafinesque (1818) created the name, Criosanthes borealis, for the same species and, in 1837, enriched his new genus with the taxon Criosanthes parviflorum. Even the smallest differences, which can occasionally occur in one individual, were enough for Rafinesque to introduce a new species and even a genus into the literature.
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In the eighteen-eighties, a Frenchman, A. Franchet, received a small lady's-slipper from China, which fascinated him by the free-standing lateral sepals. In 1885 he described the plant as Cypripedium plectrochilum. Later on it was cited as C. plectrochilon, which was never used by Franchet. Right after he described it the first time, Franchet received a flower of C. arietinum from a cultivated specimen from Godefroy-Lebeuf, the editor of "L'orchidophile". Immediately Franchet realized the close similarity to C. plectrochilum and in 1886 he used this name as a synonym for C. arietinum and excused himself because Brown didn't talk about the free-standing sepals in his first description.
Franchet thought about putting Cypripedium arietinum into its own genus, because of the different shape of the lip, the different shaped staminodium and mainly because of the separate sepals. He did not formally publish this. In this context, he named the genus Arietinum. In 1833, Beck described C. arietinum as Arietinum americana. Criosanthes would have been the prior idea for the genus name, but obviously Franchet didn't know that name at all. Even if Luer (1975), in error, put the combination Criosanthes arietinum together crediting Franchet, this combination was only used by House in 1905.
In conclusion, I'd like to cite Atwood (1984) who
wants to re-introduce House's combination. His reason is based on a cladistical analysis in which he focuses on three primitive characters:
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lateral sepals free standing
lip with a spur
staminodium barely different from a fertile anther.
These characters are supposed to characterize Cypripedium arietinum as the most primitive lady's-slipper, even more so than the tropical genus Selenipedium. Therefore, he names two more characters that are shared by C. arietinum and Selenipedium:
very small epidermal cells of the leaves
thickened cells of the pericycel (the layer around the central cylinder of the root.)
and one more character that is shared by C. arietinum and Phragmipedium:
sepals are rolled within the bud. The last three characters actually point to special patterns of C. arietinum. In my opinion, the first three characters are not as convincing as Atwood thinks to describe a new genus.
The degree of separation of the lateral sepals varies in the other species of the genus very strongly, so that you can find in the sparrow's egg lady's-slipper, Cypripedium passerinum, individuals with completely separate lateral sepals. Cypripedium passerinum is a lady's-slipper with very small flowers as is C. arietinum. The function of the synsepalum (fused sepals) is to strengthen the lip. Maybe that is also an explanation for the strangely expanded lower bottom side of the lip. It could contribute to the stability of the lip and, therefore, a synsepalum would not be needed any longer. Certainly
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the bottom side of the lip is not a spur as far as the biology of the flowers is converned, because the flower of C. arietinum is an insect trap for the same pollinators as the small white lady's-slipper, C. candidum and C. parviflorum var. pubescens. The pollinators are two species of small bees of the families Halictidae and Andrenidae (Catling, 1984). Also, it can not be an original spur without function, because, referring to evolutionary knowledge, it then would have been degraded.
The staminodium of Cypripedium arietinum does have an external structure that strongly reminds one of the structure of an original anther. It is not such a primitive form like the very original C. subtropicum or the genus Selenipedium. In contrast, the staminodium is rather big in relation to the whole flower and completely developed as an attraction signal and landing place for pollinating insects. This is even clearer in the closely related C. plectrochilum.
There also is no biochemical argument to separate Cypripedium arietinum from the genus Cypripedium. The American botanist, Dr. Martha Case (1994), examined the isoenzymes of some North American Cypripedium species. She found that the isoenzymes of C. arietinum and C. parviflorum var. pubescens are more similar than those of the latter one and the pink lady's-slipper, C. acaule. Back to the limestone barrens
Meanwhile we were in an area, which had plenty of large and small groups of Cypripedium parviflorum var.
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pubescens in flower and barely frost-damaged. While searching though the surrounding Thuja woods, we finally found the first C. arietinum. The plants were in the complete shade, very small and without flowers. There was barely a difference from the sterile broad-leaved helleborine, Epipactis helleborine, which also grow in the shade. Finally, we found some flowering C. arietinum in the margins of the woods, still under trees, but they were partly exposed to the sun. They were 20-25 cm high with flowers of 3 cm on average. Some of the plants were only 2 m away from the flowering plants of C. parviflorum var. pubescens.
The flower of Cypripedium arietinum only lasts for about one week. Once pollinated, the upper sepal comes down onto the opening of the slipper within one day. All of the 16 flowering ram's-head lady's-slippers we found on July 4, 1993, showed completely developed fresh flowers. Thus it must be the main flowering time for this area. The black-brownish soil had a pH of 5.87 (measured with 0.1 n KOH, pH 6.44 with H20) and a humus content of 78%. The water content is 62%.
We were not only there for observation, but also
we wanted to collect some material for comparative isoenzyme analyses of Cypripedium arietinum and C. plectrochilum for Martha Case. Therefore, we took only one leaf of each plant selected. We collected 34 leaves from the 80 plants so that we would get a representative average of the population.
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In the late afternoon we were at the parking lot on the shore of Lake Ontario so as not to be bothered by the annoying insects. There we cut the leaves into halves, rolled them up and put them each into a plastic tube with a cover. Then everything was put into a Styrofoam box with ice. In the morning, we took leaves of Cypripedium plectrochilum that were cultivated in the museum in Albany and put them into the iced box. We were able to send the box to Martha Case at Michigan State University before the post office in Watertown, NY closed.
A few days later I visited another location of Cypripedium arietinum in a swampy area in the middle of the state. Here the plants are very rare. They grow on the hummocks above the swampy ground. The hummocks are elevations (around 1 m) that were built up by trunks, roots etc. The plants in the swamp don't occur in very moist or wet areas, unlike the northern small yellow lady's-slipper, C. parviflorum var. makasin (previously known only as C. parviflorum) which grows here as well.
The third, and most beautiful, location I got to know at the end of my visit to the state of New York was located on the western shore of Lake Champlain, only half an hour drive from the Canadian border. The substrate consists of marble, partly established as high cliffs above the lake. We passed cottages for miles - they were built close to the lake and have probably destroyed the habitat for many Cypripedium plants. Here you only find C. acaule and C. parviflorum var. pubescens. However,
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on private property on a peninsula of the lake, you can find a large colony of C. arietinum that is protected by the owner. Chuck was allowed to have a look once. 'I'oday, he wanted to show me a colony on a steep slope. Therefore, we drove to the end of the slope and walked on the railroad track that lead along the shore. Occasionally there were deep, narrow gorges cut into the rocks for the railroad. I preferred to listen on the rail before walking into the gorges, who knows...?
There was a specific species of rattlesnake (timber rattlesnake ed.) in this area that I would have liked to see once, as a biologist. But I suppressed this wish while I was partly forced to climb on the steep cliffs. I did not feel well anyway, because I am scared of heights and did not at all want to imagine touching such a poisonous snake while seeking for holds on the cliff. I felt much better after standing on solid ground again. Then I could be impressed by the wonderful habitat. On the rock ledges pines grow (eastern white pine, Pinus strobus, and red pine, Pinus resinosa). The ledges collected bark and litter from dead needles. In the litter thick patches of common polypody fern, Polypodium virginianum, and Cypripedium acaule, with its gorgeous flowers, grew. The substrate here is extremely acid (pH 3.12) and less nutrient-rich. Cypripedium arietinum was only found in a location where the soil was enriched with more minerals and was deeper. Besides Thuja occidentalis, there were also found sugar maple, Acer saccharum, and other deciduous trees. The soil had a pH of 4.85 (measured with 0.1 n KOH, pH 6.42 with H20). The humus content was 59%, the water content, 62%.
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Cypripedium arietinum grows in moderately acid to
neutral soils in shade or partial shade. In full shade it does not form flowers and stays small, like most of the other Cypripedium species. The soil is usually well drained. In the US, C. arietinum has its largest populations in sandy soils with limestone substrate in the upper Great Lakes of central North America (Fred Case, 1987). It can stand periodic drought better than constant wetness. The range of distribution is spread from Manitoba (Canada) over the northeast of the US from the Great Lakes to Nova Scotia (Canada).
The southwestern Chinese Cypripedium plectrochilum is very similar to C. arietinum. Its habitats are also on limestone substrate. It is found in northern Yunnan and in Sechuan in southwest China. There it grows at elevations of 2200-3000 m (Handel-Mazetti, 1936) in less shady pine woods and open areas. Looking at descriptions of habitats and mapped habitats, it seems to prefer more open habitats than C. arietinum. Phillip Cribb found C. plectrochilum at Yulongxue Shan (mountain range) in Yunnan growing between limestone rocks in a dark humus soil with pine needle litter. This soil should be quite similar to those I found in northeastern North America.
To distinguish species, Schlechter (1919) allowed Cypripedium plectrochilum to be a species. His reasons were: the less inflated labellum (in contrast to C. arietinum) which is more expanded to the front; and the much wider and bigger staminodium, which is more
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expanded to the outside and less carinated in the inside. I also can see consistent color differences. Whereas the bottom part of the lip of C. arietinum is strongly red colored with a faint reticulated pattern and contrasting to the white surface, the lip of C. plectrochilum seems to be much lighter. There is only one, in comparison, slightly dark red pattern of vertical stripes, which goes down to the lip opening. The staminodium of C. arietinum is mainly tan to olive colored; that of C. plectrochilum is white with a strong red at the tip.
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Cypripedium arietinum ram's-head lady's-slipper Windsor County, Vermont
photos by P. M. Brown 41
Cypripedium arietinum ram's-head lady's-slipper Windsor County, Vermont
photos by P. M. Brown 41
Cypripedium arietinum ram's-head lady's-slipper Windsor County, Vermont photos by P. M. Brown 41
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top:
Cypripedium plectrochilum China photo by Phillip Cribb bottom:
Cypripedium plectrochilum in hort.
photo by Charles J. Sheviak
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Because there are distinct differences between both taxa, which allow a separation into two species, I've not yet found characters that would justify putting together both species into Cypripedium arietinum. In this context, the result of the isoenzyme analysis by Martha Case will be interesting. Literature Cited: Atwood, J.T. 1984. The relationships of the slipper orchids.
Selbyana 7:129-247. Beck, L.C. 1833. Arietinum americanum. Botany of the Northern and
Middle States. p. 352 Brown, R.1813.Cypripedium arietinum in Aiton, Hortus Kewensis, ed.
2, 5:222. Catling, P M. 1984. Distribution and pollination biology of
Canadian orchids in Proceedings of the 11th. World Orchid Conference-Miami. pp.121-131.
Case, F. 1987: Orchids of the western Great Lakes region. pp.73-75. Cranbrook Institute, Bloomfield Hill, MI.
Case, M. 1994, Extensive variation in the levels of genetic diversity and degree of relatedness among five species of Cypripedium. American Journal of Botany. 81 (2): in press.
Franchet, A. 1885. Cypripedium plectrochilum. Bulletin la Societe Botanique de France, T. 32, II Serie, 7:27.
Handel-Mazzet, H. 1936. Symbolae Sinicae, Teil VII. pp.1323-1324. House. H. 1905.Criosanthes arietina. Bulletin of the Torrey Botanical
Club. 32:374. Luer, C.1975. The native orchids of the United States and Canada.
New York Botanical Garden, Bronx, NY Rafinesque, C.S.1818. Criosanthes borealis. American Monthly
Magazine and Critical Review. 2:268.
- 1837. Criosanthes parviflorum. Flora Tellularia 4:46. Philadelphia
Schlechter, R. 1919. Orchideologia Sino-japonicae Prodomus. Fedes Rep. Sp. Nov. Beihefte 4. p. 84.
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Holger Perner, Reddarallee3, D-214834 Lütau, Germany Dr. Perner writes extensively on Cypripedium and recently contributed the horticultural section to Phillip Cribb's The Genus Cypripedium (reviewed in NANOJ Sept. 1997) This article originally appeared in Die Orchidee in German and in a different format. The Journal wishes to thank Ann Holmer and Stan Folsom for their valuable assistance in translating this article, and Chuck Sheviak and Phil Cribb for the loan of slides of Cypripedium plectrochilum.
Brown: NEW TAXA & COMBINATIONS
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NEW TAXA & COMBINATIONS
Paul Martin Brown Two new taxa and three new combinations are proposed. The new taxa both represent white-flowered forms, or those lacking in anthocyanins within the flowers. Cypripedium kentuckiense C. F. Reed forma pricei P.M. Brown forma nov. TYPE: United States, Arkansas, Montgomery County. May 8, 1984. (Holotype: N. A. Native Orchid Journal 4(1):47) A forma kentuckiense floribus sine rubellipigmento (sepalis et petalis albo-viridibus) labello et staminodio immaculato) differt. Differing from the forma kentuckiense in its flowers lacking all reddish pigmentation: sepals and petals clear green, lip pure white without any markings; staminode yellow without markings. Typical plants of Cypripedium kentuckiense vary from ivory-white to yellow lips but always with petals and sepals marked in some degree with dark reddish/purple pigmentation. Often this pigmentation
Brown: NEW TAXA & COMBINATIONS
46
is also present on the underside of the lip and the staminode. Named for its discoverer, Jack Price, of Blanchard, Louisiana, who first brought this most unusual form to my attention. Jack tells of first finding the plants on a hillside in southern Arkansas near Collier Spring. He and his wife had been botanizing for yellow-flowered lady's-slippers that day and had encountered great numbers. At this point Jack ventured uphill and found this colony. There were seven flowering plants present with several others in bud that may also have been this form. A similar situation, with all pigmentation lacking, occurred in Cypripedium montanum forma praetertinctum Sheviak as described in Rhodora 92:47-49. 1990. Eulophia alta Fawcett & Rendle forma pelchatii P.M. Brown forma nov. TYPE: United States, Florida, Collier County. 3 Nov. 1996. (Holotype: N. A. Native Orchid Journal 4(1):48.) Forma floribus albido-viridis conspecibus diversa. Differing from the species by its white and green flowers.
Brown: NEW TAXA & COMBINATIONS
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Cypripedium kentuckiense forma pricei ivory-lipped lady's-slipper, white-flowered form Montgomery Co., Arkansas
photos by Jack Price 47
Brown: NEW TAXA & COMBINATIONS
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top:
Eulophia alta wild coco typical coloration bottom:
Eulophia alta forma pelchatii green petals and sepals with white lip Collier Co., Florida photos by Cliff Pelchat
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Cliff Pelchat writes of the discovery:
There are about 200 species of orchids both terrestrial and epiphytic in the genus Eulophia and most of these are found in Africa. In Florida, Eulophia alta, commonly called wild coco, is a locally common terrestrial orchid found in most of the central and southern counties. We have observed it growing in Brevard and Collier counties. It has a reported range in the Western Hemisphere from Central Florida to South Florida, Mexico, the West Indian Islands, Central America and South America.
All of the plants we have observed from Brevard county south into Collier County have the typical color form as described in Carlyle Luer's book on native Florida orchids (1972). They typically have bronze colored petals and sepals with bronze to red and sometimes paler colored lips. At times they have been reported as having green sepals and petals with some pale colored forms of the lip. There are pictures (plate 72) in Luer's book illustrating two of the pale colored forms. These are known as Eulophia alta forma pallida (Brown, 1995). One picture shows a flower with green sepals and petals and the lip has a white center with a fringe of red.
We found a single plant in the Big Cypress Swamp Preserve on November 3, 1996 that is a white flowered form of Eulophia alta. It was growing near our campsite in a patch of woods that most likely had been disturbed by clearing carried out to create primitive campsites for hunters. The patch was approximately 60 feet long by 30 feet wide with a couple of slash pine trees and some palmetto bushes. In this same patch of woods we counted 14 plants, 3 in fruit with waning leaves, one in bud, and the white flowered form plant in full bloom. The plant in bud was 60 feet from the one in bloom and the buds showed potential for being white flowered also because there was a lack of the darker colors in the sepals. Most of the other plants had already set capsules that were well on their way to ripening. The white flowered plant, which was growing close to the base of a
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Palmetto bush, had 18 flowers with all white lips, green sepals and petals, and not a trace of red. Needless to say we were very excited to find this plant because all of the Eulophia alta we have observed from Brevard County south have been red or some form of red color. On Bear Island we have found numerous plants of the red color form growing in the pinewoods usually close to pine trees. John Beckner, of the Marie Selby Botanical Gardens, has told me that this color form has been observed in the past and Ken Roberts has told me that Ken Anderson of Bradenton, Florida has white-flowered Eulophia alta in cultivation. Other than these reports I know of no other documented observation of this form growing in the wild, and, based on our observations I would be confident in stating that this was a rare find.
We have kept a close observation on this particular plant visiting it each time we've been in the area, On Thanksgiving weekend we visited the site to find that 13 of the 18 flowers had been successfully pollinated and there were ripening seed capsules. On February 23, 1997 the same plant had three capsules left at the very top of the raceme which looked to be about 2 -3 weeks away from full maturity. These three capsules were green and healthy, but there was no trace of any of the other capsules. Most of the other plants in the area had completely disappeared due to the deciduous nature of this species even though there were some racemes with seed capsules. Many of the seed capsules on other plants had turned brown and did not appear to have reached maturity. An inspection of one revealed that the seed inside was black and had evidently been attacked by some form of fungus. There was also evidence of the fungus growing on the outside of the capsules in the form of a black powdery coating. I suspect that the rest of the capsules on the white flowered plant we have been observing met a similar fate. Brown, P.M. 1995. New and recently published taxa. NA Nat.
Orchid J. 1(2): 132. Luer, C. 1972. The Native Orchids Of Florida, New York Botanical
Gardens, Bronx, NewYork.
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New Combinations Recent research and revisions necessitate several new combinations. Much confusion has existed in the correct generic names for many of the Spiranthiode species. Ackerman (1995) points out that Cyclopogon is preferred to Beadlea, based on the existence of intermediate species between the two genera, with Cyclopogon having priority. Cyclopogon cranichoides forma albolabia (Brown & McCartney) P.M. Brown comb. nov. Basionym: Beadlea cranichoides forma albolabia Brown & McCartney. North American Native Orchid Journal 1(2):8. From Wes Higgins at the University of Florida whose research on the genus Encyclia has resulted in extensive new combinations at the species level, the follow new combinations are needed for our Florida species. An ongoing systematic study of the genus Encyclia based on holomorphology has determined that the genus is neither morphologically cohesive nor monophyletic. In a preliminary molecular study, analysis of the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA supports the morphological conclusion that the Encyclia subgenus Osmophytum clade should be raised to the generic level because these species are sister to the Cattleya-Laelia clade not to Encyclia subgenus Encyclia. However, the monophyly of the three currently recognized subgenera of Encyclia i.e., Encyclia subg. Osmophytum, Encyclia subg. Encyclia, and
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Encyclia subg. Dinema, is supported by cladistic analysis of both morphological and molecular data. Encyclia subgenus Osmophytum is raised to generic level and treated as Prosthechea.
Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins comb. nov. Basionym: Epidendrum erythronioides Small, Fl. South-eastern US. 328, 1329. 1903. Synonym: Encyclia boothiana (Linnaeus) Dressler var. erythronioides (Small) Luer. Luer, Nat. Orchids of Fl. 204. 1972. Prosthechea cochleata (Linnaeus) W. E. Higgins var. triandra (Ames) W.E. Higgins comb. nov. Basionym: Epidendrum cochleatum Linnaeus var. triandrum Ames, Contrib. Ames Bot. Lab. 1:16. 1904. Synonym: Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler. Luer, Nat. Orchids of Fl. 202. 1972. Prosthechea cochleata (Linnaeus) W. E. Higgins var. triandra (Ames) W.E. Higgins forma albidoflava (P. M. Brown) P. M. Brown comb. nov. Basionym: Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler forma albidoflava P.M. Brown. Brown, P.M. NA Native Orchid Journal 1(2): 131. 1995.
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In addition to general membership information it contains a full set of Tables of Contents for all issues - Volume 1, 1995-Volume 3, 1997 - and will be updated as each issue is published. Other information of interest and occasional articles will also be available on-line. Special thanks to the Florida Museum of Natural History at the University of Florida for hosting our website. We appreciate the assistance of Dick Ruble in the FLMNH Office of Museum Technology and Kent Perkins, Manager of the Collection, in the FLMNH/University of Florida Herbarium.
Empiricist: SETTING GOALS
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SETTING GOALS
The Slow Empiricist
As spring slowly advances up the Northern Hemisphere, awakening the dormant flora with its gentle caress, the botanical enthusiast is also awakening from his/her long winter's nap. Of course this is not so much the case for those fortunate enough to live in the more southerly regions, or for those who can afford a mid-winter vacation to the south, where there is still evidence of things growing and blooming all season. For many, spring signals the beginning of another season of botanizing. This article aims at helping you to prioritize your activities and achieve a satisfying set of experiences for the 1998 calendar year. If you are like me, you probably try to crowd too much into your schedule and end up frustrated when you can't meet all of your plans fully. A calendar comes in very handy as you start to map out your year's activity. This way you can see what days and weeks are available to you for your intended activities. If you want to take a course, or plan an extended field excursion (if you know the approximate blooming times of the orchids you want to see), calendars will help you choose the correct dates for your activities. Once you have identified your needs
Empiricist: SETTING GOALS
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and wants, you have a variety of ways to organize yourself to meet those needs. I am going to deal mostly with ways to successfully plan your field experiences because once you have identified the courses you would like to enroll in for the coming season, you have most of the planning already done for you by the schedule of the course. If you read the rest of this article, however, the attitudes that I am trying to foster may help you get more out of the instruction you are receiving. The underlying principles that apply to fieldwork should have some carry-over for you in your course work.
Different people approach tasks in different ways. Some like to be totally organized with every minute accounted for. On the other extreme, there are individuals who hate to be constrained by schedules and rigid boundaries. The totally organized are like the tourists who visit a foreign country and have spent the entire year before their trip reading up on the place. They have set a schedule of visits to museums, galleries and note-worthy landmarks that cannot be easily breached. The free-spirited individual has a general idea of what is in the foreign country but cannot bring himself/herself to plan any kind of itinerary to follow. In both cases, they may miss important experiences: one, because their rigid schedule won't permit it; the other, because they hadn't done enough homework to be aware of what they were missing.
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For my own self, I like to have an underlying structure to base my plans on that allows me the freedom to take advantage of the unexpected occurrence. This approach avoids the pitfalls of being too lax or too tied to a goal. To me, you should always be open to the unexpected, weighing the possibilities and making a decision that will best answer the situation for you and your companions. I would like to go on record as saying that I don't always follow this advice I have freely shared with you, the reader. I have become so involved with my activity that time slips beyond my awareness and I have found myself far afield, enticed by the plants I am finding. Ruefully, I find myself running a race with the fleeting daylight to regain civilization.1 Usually, the next time I am botanizing I remember the last experience and don't over-extend myself.
Whatever type of organizer you are, you might be
more successful if you set down the goals you would like to attain this year on a sheet of paper. A list made on a computer is even better because you can rearrange the items with simple maneuvers as you think through what you really want to do with your time. So once you have set your activities in some kind of order of importance, it would be wise to look at the incidentals that can affect the success or failure of your quest.
For many of you, I suspect, the top priority is to locate a new member of the Orchidaceae that has eluded you in other years. This may entail travel, which may
1 See Flops and Failures, NANOJ 2(4): 350-360. 1996.
Empiricist: SETTING GOALS
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require time, and other resources like monetary expenses as well as information as to the whereabouts of said plants. It may also encompass traveling companions who have to be considered. If you are using your computer to help set your goals, you can insert sections that cover these areas to help you see what is required of you in order to meet those goals. If you are more into making hand-written lists you might be wise to create a list in a loose leaf-leaf or spiral-bound notebook. If you put a heading at the top of each page of what you want to accomplish, you can add the things that need to be done to achieve your goal under the appropriate heading. I would suggest that you determine the order of importance of your goals for 1998. Then you can avoid over-crowding your schedule by having specific areas that you have set up in order of importance. By doing this kind of list-making you can weed out those things that take undue amounts of your time and attention and allowing the really important things to be concentrated on to become richer and fuller experiences.
This writing down of your goals may sound
daunting to the free-spirited individual, so I would suggest that if you are inclined not to be constrained by rigid plans, you might hold in your mind the following things as you lay your plans. The following suggestions are also important headings for the highly organized individual to include in his/her lists of things to take into consideration. A successful quest in the field will require an appropriate amount of time to complete it, so you should think about the time requirements. Location is another need that requires attention. Have you got
Empiricist: SETTING GOALS
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accurate information and good maps to guide you? Will your quest require an overnight stay? This can be especially true for distant sites that require four or more hours to reach. Traveling eight or more hours in one day, combined with fieldwork, usually makes the return trip very tiring. If this is a new site for you, it will probably take more time than you thought to find your quarry. It might be wise to set an early departure time to allow for exploration, or plan for an overnight near the site.
Finally, think of the needs that are important to
you. Are you a photographer? What do you need to include in your equipment to insure a successful session with the plants you are hoping to photograph? What about the weather? Have you included the appropriate clothing to give you comfort as you explore the territory? How far will you be from facilities that you require like bathrooms or restaurants? Keeping an awareness of your expectations will help you better meet them when you are in the actual situation.
If you do a little preparation before you venture
into new areas, the success rate for your adventure should be higher. You still need to be open to unexpected events that come across your path. I have seen single-minded individuals pass up plants that they have come upon accidentally, as they strive to reach their new enthusiasm. Conversely, I have seen people become so distracted by the unexpected that they lose sight of their intended goal. For me, I try to keep things in some kind of balance. If I consider the accidental discovery,
Empiricist: SETTING GOALS
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appreciate it for what it is, and weigh it against my goal for that day, I can make a reasonable judgement of what should be done. If I find that the quest to find that elusive orchid I have never seen before is becoming too complicated (by the new discovery), or if the weather, distance, or some other constraint is affecting my enjoyment, I can give up the attempt of reaching my goal. Although it has taken me a long time to realize the fact, I have come to know that by not succeeding, I will still have the goal to try for on another day or in another year.2 I can also rationalize that, although frustrated, all is not lost; I have not done my homework in vain for I have grown in knowledge and experience.
To summarize: If you can prepare yourself for the
coming season of exploration and grow in your understanding of what you need to put into such activities you will have a more successful time. Whether you keep detailed lists or just have a general idea in the back of your head as to what is required, by being aware of the possibilities, you should have a more rewarding year of botanizing. I approach my botanical work by using a little of both methods. That way, I can have what to me is the best of both worlds. Before I go out into the field to explore, I can be prepared with the correct information as to location and times of blooming. I try to know as well what I have to put into the effort as far as travel, expense and the physical effort needed to meet my goal while I still remain flexible enough to take
2 See Flops and Failures, NANOJ 2(4): 350-360. 1996.
Empiricist: SETTING GOALS
60
advantage of the unexpected that I may encounter. Lastly, I try not to view a failed objective as something negative, but something I can learn from and, hopefully, grow from, to be more competent in the future.
The Slow Empiricist
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61
CHECKLIST OF THE ORCHIDS OF NORTH
AMERICA north of Mexico
Since the publication of Carlyle A. Luer's two volume
work, (1972, 1975) on the orchids of the United States and
Canada much taxonomic work has taken place. Schrenk's (1977)
checklist was reasonably comprehensive, but in addition to being
generally unavailable to most orchidists, it contained some very
different concepts of genera and species. The 2nd edition of
Kartesz' (1994) Checklist of US and Canadian plants brought a
more realistic list of species, but did not deal with several newer
species, genera concepts or taxa at the forma level. This checklist
has been compiled in order to help coordinate all current North
American orchid taxonomic information. Literature citations are
given for recently described, little known or significantly revised
taxa.
Synonyms are given if the names differ from those used by
Luer.
Coleman (1995) should be consulted for the most current
information on the genus Piperia.
Garay (1982) and Catling (1989) are followed for most of the
generic changes in Spiranthes, although Ackerman (1995) is
applicable to all of the Florida taxa.
Sheviak's recent work on the green-flowered Platanthera has
helped considerably in sorting out these confusing taxa.
Wes Higgins' work on the genus Encyclia has resulted in
several new generic combinations.
The recent work for volume 26 of the Flora of North America,
which will contain the Orchidaceae has resulted in numerous
CHECKLIST
62
nomenclatural changes. Those changes appear in this edition of
the checklist. The publication of that volume is scheduled for
late in 2001.
White-flowered forms, often referred to as albinos, exist for
many of our native species.
Ranges given are for reference only and do not imply precise
distribution or extant populations.
Abbreviations for the states and provinces are standard and the
following should be noted: CtA=Central America; Mx=Mexico;
BA= Bahama Archipelago; WtI=West Indies; SA=South
America; RM=Rocky Mountains; AM=Appalachian Mountains;
* indicates species documented to be either introduced or
adventive
note: Chuck McCartney has recently called my attention to the
fact that several of the rarer 'lost' species in southern Florida may
represent waifs as the result of the efforts of past orchidists to
establish orchid gardens within the wild in tropical Florida. This
could easily account for the one-time records for several of the
more southerly species. One possible example could be Pelexia
adnata.
References:
Ackerman, J. D. 1995. An Orchid Flora of Puerto Rico and the
Virgin Islands. Memoirs Volume 73. New York Botanical
Garden, Bronx, NY.
Catling, P.M. 1989. Biology of North American representatives
of the subfamily Spiranthoideae, in North American Native
Terrestrial Orchid Propagation and Production. Brandywine
Conservancy. Chadds Ford, PA.
Coleman, R.A. 1995. The Wild Orchids of California. Piperia:
105-135. Comstock Publishing Associates/Cornell University
Press, Ithaca, NY.
Garay, L.A. 1982. A generic revision of the Spiranthinae,
Botanical Museum Leaflet, Harvard University 28: (4): 277-425.
CHECKLIST
63
Hammer, R. L. 1981. Finding New Orchids, a Contribution to
the Orchidaceae of Florida. Fairchild Tropical Garden Bulletin
36(3): 16-18.
1992. The Strange Case of Carter's Orchid.
Fairchild Tropical Garden Bulletin 47(2): 34-39.
2001. Status report on the orchids of southern
Florida. North American Native Orchid Journal 7(1)
Kartesz, J.T. 1994. A synonymized checklist of the vascular
plants of United States, Canada and Greenland. 2nd. ed. Timber
Press. 2 vols.
Luer, Carlyle A. 1972. The native orchids of Florida. New York
Botanical Garden, Bronx, NY
1975. The native orchids of the United States &
Canada excluding Florida. New York Botanical Garden, Bronx,
NY
McCartney, C.L., Jr. 1981. The orchids of Rabenau Camp -- 1.
The Epiphytes. American Orchid Society Bulletin 50(5): 527-
535.
1981. The orchids of Rabenau Camp -- 2. The
Terrestrials. American Orchid Society Bulletin 50(6): 653-660.
1985. The Orchids of Everglades National
Park -- 1. American Orchid Society Bulletin 54(3): 265-276.
1987. A thrice annotated checklist of the orchids
of southeastern Florida. (Revised edition.) Published privately.
2226 Lincoln St., Apt. 3, Hollywood, FL 33020.
1992. Orchids of south Florida's rock pinelands.
Fairchild Tropical Garden Bulletin 47(2): 12-33.
Schrenk, W.J. 1977. Zussammenstellung der Orchideenarten der
Vereinigten Staaten von Amerika und der amerikanishen
Jungferninsein. Die Orchidee 28: 98-104.
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64
Amerorchis rotundifolia (Banks) Hulten
SMALL ROUND-LEAVED ORCHIS
AK - NF s to MT, WY; MN - ME
forma angustifolia Rousseau - narrow-leaved form
beckettiae (Boivin) Hulten - white-flowered form
immaculata Mazurski & L.P. Johnson - white-lipped form
lineata (Mousley) Hulten - lined-lip form
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 132.
Johnson, L.P. 1995. Lindleyana 10(1): 1.
Aplectrum hyemale (Mulhenberg ex Willdenow) Torrey
PUTTY-ROOT; ADAM AND EVE
MN - MA s to AR - GA
forma pallidum House - yellow-flowered form
Arethusa bulbosa Linnaeus
DRAGON'S-MOUTH
MN - NF s to NC
forma albiflora Rand & Redfield - white-flowered form
subcaerulea Rand & Redfield - lilac-blue flowered form
Basiphyllaea corallicola (Small) Ames
CARTER'S ORCHID
seFL; BA, WtI
Hammer, R. L. 1992. Fairchild Trop. Gard. Bulletin. 47(2): 34-39.
McCartney, C.L., Jr. 1991 . Florida Orchidist. 34(4): 136-157.
_______________ 1992. Florida Orchidist. 35(1): 195-211.
Beloglottis costaricensis (Reichenbach f.) Schlechter
SYN: Spiranthes costaricensis Reichenbach f.
COSTA RICAN LADIES'-TRESSES
sFL; CtA, nSA
Bletia patula Hooker
HAITIAN PINE-PINK*
se FL; WtI
Bletia purpurea (Lambert) de Candolle
PINE-PINK
forma alba (Ariza-Julia & J. Jiménez Alm.) P.M. Brown - white-flowered
form
sFL; WtI, CtA, nSA
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65
Brown, P.M. 2000. NA Native Orchid Journal 6(4):
Bletilla striata (Thunberg) Reichenbach f.
URN ORCHID*
wFL
Brassia caudata (Linnaeus) Lindley
SPIDER ORCHID
sFL; WtI, CtA, nSA
Bulbophyllum pachyrhachis (A. Richard) Grisebach
RAT-TAIL ORCHID
sFL; WtI, CtA, SA
Calopogon barbatus (Walter) Ames
BEARDED GRASS-PINK
NC - FL w to LA
Calopogon multiflorus Lindley
MANY-FLOWERED GRASS-PINK
GA - FL w to MS
Calopogon oklahomensis D.H. Goldman
OKLAHOMA GRASS-PINK
s. IN, eKS, eOK, swMO, AK, eTX, wcLA
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 133.
Goldman, D.H. 1995. Lindleyana 10(1): 37-42.
Goldman, D.H. and S. Orzell. 2000. Lindleyana 15(4): 237-251.
Calopogon pallidus Chapman
PALE GRASS-PINK
VA - FL w to LA
forma albiflorus P.M. Brown - white flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 8.
Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var.
tuberosus
GRASS-PINK
[including var. latifolius (St. John) Fernald]
MN - NF s to FL w to TX
forma albiflorus Britton - white flowered form
Catling, P.M. & Z. Lucas. 1987. Rhodora 89: 401-413.
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66
Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg var.
simpsonii (Small) Magrath
SIMPSON'S GRASS-PINK
sFL
forma niveus P.M. Brown - white-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 130.
Magrath, L.K. & J.L. Norman. 1989. SIDA 13(3): 371-372.
Calypso bulbosa (Linnaeus) Oakes var. americana (R.Brown) Luer
EASTERN FAIRY-SLIPPER
AK - NF s in the RM; s to upper GtL, n New Eng.
forma albiflora P.M. Brown - white-flowered form
rosea P.M. Brown - pink-flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 17.
Calypso bulbosa (Linnaeus) Oakes var. occidentalis (Holtzman)Boivin
WESTERN FAIRY-SLIPPER
AK - CA e to ID
forma nivea Brown & Coleman - white-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 17.
Coleman, R.A. 1992. AOS Bulletin 61(8): 776-781/fc.
Campylocentrum pachyrrhizum (Reichenbach f.) Rolfe
CROOKED-SPUR ORCHID
sFL; WtI, nSA
Cephelanthera austiniae (A. Gray) Heller
PHANTOM ORCHID
sBC - CA e to ID
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana 8(2): 79.
Cleistes bifaria (Fernald) Catling & Gregg
SYN: Pogonia bifaria P.M. Brown & Wunderlin
UPLAND SPREADING POGONIA
WV - FL w to eLA
Brown & Wunderlin. 1997. NA Native Orchid Journal 3(4): 450-452.
Catling, P.M. & K.B. Gregg. 1992. Lindleyana 7(2): 57-73.
Cleistes divaricata (Linnaeus) Ames
SYN: Pogonia divaricata (L.) R. Br.
SPREADING POGONIA
NJ - FL
forma leucantha P.M. Brown - white-flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 7.
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67
Catling, P.M. & K.B. Gregg. 1992. Lindleyana 7(2): 57-73.
Coeloglossum viride (Linnaeus) Hartman var. viride
NORTHERN BRACTED GREEN ORCHIS
[including var. islandicum (Lindley) Schulze and Habenaria viridis var.
interjecta Fernald]
AK - NF; Eurasia
Coeloglossum viride (Linnaeus) Hartman var. virescens (Mulhenberg) Luer
LONG BRACTED GREEN ORCHIS
AK - NF s to WA, NM, IA, NC
Corallorhiza bentleyi Freudenstein
BENTLEY'S CORALROOT
Bentley, S. 2000. Native Orchids of the Southern Appalachians, pp. 71-75.
Freudenstein, J.V. 1999. Novon 9(4): 511-513.
Corallorhiza maculata (Rafinesque) Rafinesque var. maculata
SPOTTED CORALROOT
BC - NF s to CA, AZ, NM; AM to nGA
forma flavida (Peck) Farwell - yellow-stemmed form
rubra P.M. Brown - red-stemmed form
Brown, P.M..1995. NA Native Orchid Journal 1(1): 8.
Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley)
Ames
WESTERN SPOTTED CORALROOT
BC - NF s to CA, AZ, NM, MN, New Eng., VA
forma aurea P.M. Brown - golden yellow/spotted form
immaculata (Peck) Howell - yellow spotless form
intermedia Farwell - brown-stemmed form
punicea (Barth.) Weatherby & Adams - red-stemmed form
Brown, P.M.. 1995. NA Native Orchid Journal 1(3): 195.
Freudenstein, J.V. 1986. Contr. Univ. Mich. Herb. 16: 145-153.
1997. Harvard Papers in Botany 10:5-51.
Corallorhiza mertensiana Bongard
WESTERN CORALROOT
sAK - CA e to MT, wWY
forma albolabia P.M. Brown - white-flowered form
pallida P.M. Brown - pale colored form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 9.
Brown, P.M.. 1995. NA Native Orchid Journal 1(3): 197.
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Corallorhiza odontorhiza (Willdenow) Nuttall var. odontorhiza
AUTUMN CORALROOT
SD - ME s to OK - GA
forma flavida Wherry - yellow-stemmed form
Corallorhiza odontorhiza (Willdenow) Nuttall var. pringlei (Greenman)
Freudenstein
PRINGLE'S AUTUMN CORALROOT
IA - ME s to TN, NC; MX
Freudenstein, J.V. 1993. Dissertation. Cornell University.
1997. Harvard Papers in Botany 10:5-51.
Corallorhiza striata Lindley var. striata
STRIPED CORALROOT
BC - PQ s to CA, NY
forma eburnea P.M. Brown - yellow/white form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 9.
Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams
SYN: Corallorhiza striata forma fulva Fernald
VREELAND'S STRIPED CORALROOT
CA - ND s to NM; PQ; MX
forma flavida (Todson & Todson) P. M. Brown -
yellow/white form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 14.
Corallorhiza trifida Chatelain
EARLY CORALROOT
[including var. verna Fernald]
AK - NF s to OR, s in RM e toWV; Eurasia
Corallorhiza wisteriana Conrad
WISTER'S CORALROOT
MT - NJ s to AZ - FL; MX
forma albolabia P.M. Brown - white-flowered form
rubra P.M. Brown ined. - red-stemmed form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 9-10.
Cranichis muscosa Swartz
MOSS-LOVING CRANICHIS
sFL; WtI, CtA, nSA
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Cyclopogon cranichoides (Grisebach) Schlecter
SYN: Beadlea cranichoides (Grisebach) Small
Spiranthes cranichoides (Griesebach) Cogniaux
SPECKLED LADIES'-TRESSES
FL; BA, WtI, CtA, SA
forma albolabia (Brown & McCartney) P.M. Brown - white-lipped form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 8.
1998. NA Native Orchid Journal 4(1):52
Cyclopogon elatus (Swartz) Schlecter
SYN: Beadlea elata (Swartz) Small
Spiranthes elata (Swartz) L.C. Richard
TALL NEOTTIA
sFL; WtI, MX, CtA, nSA
Cypripedium acaule Aiton
PINK LADY'S-SLIPPER; MOCCASIN FLOWER
NWT - NF s to MN - GA
forma albiflorum Rand & Redfield - white-flowered form
biflorum P.M. Brown - two-flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(3): 197.
Cypripedium arietinum R. Brown
RAM'S-HEAD LADY'S-SLIPPER
MAN - NS s to MN - MA
forma albiflorum House - white-flowered form
biflorum P.M. Brown - two-flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(3): 198.
Cypripedium californicum Gray
CALIFORNIA LADY'S-SLIPPER
OR-CA
Cypripedium candidum Mulhenberg ex Willdenow
SMALL WHITE LADY'S-SLIPPER
SAS - MN - wNY s to MO, KY, NJ
Cypripedium fasciculatum Kellogg ex S. Watson
CLUSTERED LADY'S-SLIPPER
WA - CA e to MT s to CO
Brownell , V.R. & P.M. Catling. 1987. Lindleyana 2(1): 53-57.
Elliman, T. & A. Dalton. 1995. NA Native Orchid Journal 1(1): 59-73.
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Cypripedium guttatum Swartz
SPOTTED LADY'S-SLIPPER
AK - NWT; Siberia
Cypripedium kentuckiense C.F. Reed
KENTUCKY LADY'S-SLIPPER
neVA; KY - eTX
forma pricei - white-flowered form
Atwood, J. T. Jr. 1984. AOS Bulletin 53(8): 835-841.
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 255.
1998. NA Native Orchid Journal 4(1): 45.
Reed, C. 1981. Phytologia 48(5): 426-428.
Weldy, T. W., H. T. Mlodozeniec, L. E. Wallace & M. A. Case. 1996.
Sida 17(2): 423-435.
Cypripedium montanum Douglas ex Lindley
MOUNTAIN LADY'S-SLIPPER
sAK - CA e to ALB-WY
forma praetertinctum Sheviak - white-petaled form
welchii P.M. Brown - crimson edge-lipped form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 198.
Sheviak, C.J.. 1990. Rhodora 92: 47-49.
Cypripedium parviflorum Salisbury var. parviflorum
SYN: Cypripedium calceolus Linnaeus var. parviflorum Salisbury
SOUTHERN SMALL YELLOW LADY'S-SLIPPER
KS -MA s to LA - GA
forma albolabium Magrath & Norman - white-lipped form
Magrath, L.K. & J.L. Norman. 1989. SIDA 13(3): 371-372.
Sheviak, C.J. 1994. AOS Bulletin 63(6): 664-669.
__________. 1995. AOS Bulletin 64(6): 606-612.
__________. 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak
SYN: Cypripedium calceolus Linnaeus var. parviflorum Salisbury
NORTHERN SMALL YELLOW LADY'S-SLIPPER
BC - nCA - NF s to IL - PA
Sheviak, C.J. 1993. AOS Bulletin 62(4): 403.
__________. 1994. AOS Bulletin 63(6): 664-669.
__________. 1995. AOS Bulletin 64(6): 606-612.
__________. 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight
SYN: Cypripedium calceolus Linnaeus var. pubescens (Willdenow) Correll
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71
LARGE YELLOW LADY'S-SLIPPER
(including var. planipetalum Fernald)
AK - NF s to AZ - GA
Sheviak, C.J. 1994. AOS Bulletin 63(6): 664-669.
__________.1995. AOS Bulletin 64(6): 606-612.
__________. 1996. NA Native Orchid Journal 2(4): 319-343.
Cypripedium passerinum Richmond var. passerinum
SYN: Cypripedium passerinum Richmond var. minganense Victorin
SPARROW'S EGG LADY'S-SLIPPER; FRANKLIN’S LADY’S-
SLIPPER
AK - PQ s to n MT
Cypripedium reginae Walter
SHOWY LADY'S-SLIPPER
SAS -NF s to AR, n. AL
forma albolabium Fernald & Schubert - white-flowered form
Cypripedium yatabeanum Makino
SYN: Cypripedium guttatum var. yatabeanum (Makino) Hulten
YELLOW SPOTTED LADY'S-SLIPPER
AK; Siberia, n Japan
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 199.
Hybrids:
Cypripedium xalaskanum P.M. Brown
ALASKAN SPOTTED LADY'S-SLIPPER
(C. guttatum x C. yatabeanum)
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 199.
Cypripedium xandrewsii Fuller nm. andrewsii
ANDREWS' LADY'S-SLIPPER
(C. candidum x C. parviflorum var. makasin)
Cypripedium xandrewsii Fuller nm. favillianum (Curtis) Boivin
FAVILLE'S LADY'S-SLIPPER
(C. candidum x C. parviflorum var. pubescens)
Cypripedium xandrewsii nm. landonii (Garay) Boivin
LANDON'S LADY'S-SLIPPER
(C. candidum x C. xandrewsii nm. favillianum)
CHECKLIST
72
Cypripedium xcolumbianum Sheviak
COLUMBIA LADY'S-SLIPPER
(C. parviflorum x C. montanum)
Sheviak, C.J. 1992. AOS Bulletin 61(6): 546-559.
Cyrtopodium polyphyllum(Vell) Pabst ex F. Barrios
YELLOW COWHORN ORCHID*
FL; SA
in Luer, 1972 as C. andersonii (Lambert ex Andrews) R. Brown; p. 234,
pl. 71:1.
Menezes, L.C. 1995. AOS Bulletin 64(3): 248-251.
Hammer, R. 1997. NA Native Orchid Journal 3(2): 194-202.
Cyrtopodium punctatum (Linnaeus) Lindley
COWHORN ORCHID
FL; Mx, CtA, WI, SA
Dactylorhiza aristata (Fischer ex Lindley) Soo var. aristata
FISCHER'S ORCHID
AK; ne Asia
forma alba P.M. Brown - white-flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 9.
Dactylorhiza aristata (Fischer ex Lindley) Soo var. kodiakensis Luer &
Luer f.
KODIAK ORCHID
AK
forma rosea P.M. Brown - pink form
perbracteata (Lepage) P.M. Brown - leafy, flowerless form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 199, 241.
Brown, P.M. 1996. NA Native Orchid Journal 2(1): 101.
Ospina H., M. 1997. NA Native Orchid Journal 3(3): in press
Dactylorhiza cf. fuchsii (Druce) Soo
TIMMINS MARSH ORCHID*
ON; Europe
in Luer, 1975 as D. maculata (Linnaeus) Soo; p. 160, pl. 39:8-9.
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana 8(2): 80-81.
Dactylorhiza praetermissa (Druce) Soo
SYN:Dactylorhiza majalis (Reich.f.) Summerhays subsp. praetermissa
(Druce) D.M.Moore & Soo
SOUTHERN MARSH ORCHID
CHECKLIST
73
NF; Europe
Catling,P.M. & C.J. Sheviak. 1993. Lindleyana 8(2): 80-81.
Meades, S. 1994. Saraccenia. 5(1): 13-15.
________ 1995. NA Native Orchid Journal 1(3): 245.
Dactylorhiza praetermissa (Druce)Soo var. junialis (Vermeulen) Senghas
SYN: Dactylorhiza majalis (Reich.f.) Summerhays subsp. praetermissa
(Druce) D.M.Moore & Soo var. junialis (Vermeulen) Senghas
LEOPARD MARSH ORCHID
NF; Europe
Clase, H.J. & S.J. Meades. 1996. NA Native Orchid Journal 2(3): 208-
217.
Deiregyne confusa Garay
In Luer as Deiregyne durangensis (Ames & Schweinfurth) Garay
SYN: Spiranthes durangensis Ames & Schweinfurth
DURANGO LADIES'-TRESSES
TX; MX
Dendrophylax lindenii (Lindley) Bentham ex Rolfe
SYN: Polyrrhiza lindenii (Lindley) Cogniaux
Polyradicion lindenii (Lindley) Garay
GHOST ORCHID; FROG ORCHID
sFL; CUBA
Dichromanthus cinnabarinus (Llave & Lexara) Garay
SYN: Spiranthes cinnabarina (Llave & Lexara) Hemsley
CINNABAR LADIES'-TRESSES
TX; MX
Eltroplectris calcarata (Swartz) Garay & Sweet
SYN: Centrogenium setaceum (Lindley) Schlecter
SPURRED NEOTTIA
FL; BA, WtI, nSA
Encyclia rufa (Lindley) Britton & Millspaugh
REDDISH EPIDENDRUM
sFL
Encyclia tampensis (Lindley) Small
FLORIDA BUTTERFLY ORCHID
FL
forma albolabia P.M. Brown - white lipped form
CHECKLIST
74
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 131.
Epidendrum acuñae Dressler
ACUÑA'S STAR ORCHID
sFL; CtA; WtI
Epidendrum amphistomum A. Richard
DINGY-FLOWERED EPIDENDRUM
sFL; WtI, CtA, nSA
forma rubrifolium P.M. Brown - red-leaved form
Brown, P.M. 2000. NA Native Orchid Journal 6(1): 61.
Hagsater, E. 2000. NA Native Orchid Journal 6(4): 300-310.
Epidendrum floridense Hagsater
SYN: Epidendrum difforme Jacquin in part
Neolehmannia difformis (Jacquin) Pabst
FLORIDA UMBELLED EPIDENDRUM
sFL; WtI, CtA, n + cSA
Hagsater, E. 2000. NA Native Orchid Journal 6(4): 300-310.
Hagsater, E. & G. Salazar. 1993. Icones Orchidacearum
Romero, G.A.1994. A.O.S. Bulletin 63(10): 1168-1170.
Epidendrum magnoliae var. magnoliae
GREEN-FLY ORCHIS
NC s to FL w to LA
Hagsater, E. 2000. NA Native Orchid Journal 6(4): 300-310.
Epidendrum magnoliae var. mexicanum (L.O. Williams) P.M. Brown
BRONZE GREEN-FLY ORCHIS
FL; MX
Brown, P.M. 1998. NA Native Orchid Journal 5(1):3
2000. NA Native Orchid Journal 6(4): 337-338.
Epidendrum nocturnum Jacquin
NIGHT-FRAGRANT EPIDENDRUM
sFL; WtI, CtA, n + cSA
Epidendrum radicans Paven ex Lindley
CLIMBING EPIDENDRUM*
sFL
Epidendrum rigidum Jacquin
RIGID EPIDENDRUM
sFL; WtI, CtA, n + cSA
CHECKLIST
75
Epidendrum strobiliferum Swartz
CONE-BEARING EPIDENDRUM
sFL; WtI, CtA, n + cSA
Epipactis atrorubens (Hoffman ex Bernhardii) Besser
RED HELLEBORINE*
VT; Europe
Epipactis gigantea Douglas ex Hooker
STREAM ORCHID
sBC - MT s to CA, AZ, NM - SD ; MX
forma citrina P.M. Brown - yellow-flowered form
rubrifolia P.M. Brown - red-leaved form
Brown, P.M. 1995. NA Native Orchid Journal 1(4): 287.
2001. NA Native Orchid Journal 7(1) ined.
Coleman, R.A. 1995. Wild Orchids of CA. p. 75; pl. 13.
Epipactis helleborine (Linnaeus) Cranz
BROAD-LEAVED HELLEBORINE*
e US; se CAN; scattered w NA; Europe
forma alba (Webster) Boivin - white-flowered form
luteola P. M. Brown - yellow-flowered form
monotropoides (Mousley) Scoggin - albino form
variegata (Webster) Boivin - variegated form
viridens A. Gray - green-flowered form
Brown, P.M. 1996. NA Native Orchid Journal 2(4): 316.
Eulophia alta (Linnaeus) Fawcett & Rendle
WILD COCO
GA - FL; MX, WtI, CtA, SA, Africa
forma pallida P.M. Brown - pale colored form
pelchatii P.M. Brown - white flowered from
Brown, P.M. 1995 NA Native Orchid Journal 1(2): 131.
1998. NA Native Orchid Journal 4(1): 46.
Galeandra bicarinata G.A. Romero & P.M. Brown.
TWO KEELED GALEANDRA
sFL; CUBA
Romero, G.A. & P.M. Brown. 2000. NA Native Orchid Journal 6(2):
Galearis spectabilis (Linnaeus) Rafinesque
CHECKLIST
76
SHOWY ORCHIS
MN - ME s to AR - GA
forma gordinierii (House) Whiting & Catling - white-flowered form
willeyi (Seymour) P.M. Brown - pink-flowered form
Brown, P.M. 1988. Wild Flower Notes 3(1): 20.
Goodyera oblongifolia Rafinesque
GIANT RATTLESNAKE ORCHIS
sAK - NF s to ME; CA, NM; MX
forma reticulata (Boivin) P.M. Brown - reticulated leaf form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 14.
Goodyera pubescens (Willdenow) R. Brown
DOWNY RATTLESNAKE ORCHIS
ON - NS s to AR - SC
Goodyera repens (Linnaeus) R. Brown
LESSER RATTLESNAKE ORCHIS
AK - NF s to WY, sRM, sAM
forma ophioides (Fernald) P.M. Brown - white veined leaf form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 14.
Goodyera tesselata Loddiges
CHECKERED RATTLESNAKE ORCHIS
ON - NF s to MN - MD
Govenia floridana P.M. Brown
FLORIDA GOVENIA
sFL
Brown, P.M. 2000. NA Native Orchid Journal 6(3): 230-240.
in Luer, 1972 as Govenia utriculata (Swartz) Lindley; p. 243, pl.73:1-4.
Greenwood, E. W., 1991. AOS Bulletin 60(9): 867-869.
Greenwood, E.W. 1996. NA Native Orchid Journal 2(1):344-349.
Gymnadenia conopsea (Linnaeus) R. Brown
SYN: Habenaria conopsea (Linnaeus) Bentham
FRAGRANT ORCHID*
(CT)
Habenaria distans Grisebach
FALSE WATER-SPIDER ORCHID
sFL; WtI, CtA, nSA
CHECKLIST
77
Habenaria macroceratitis Willdenow
LONG-HORNED HABENARIA
FL; MX
Habenaria odontopetala Reichenbach f.
SYN: Habenaria strictissima Reichenbach f. var. odontopetala
(Reichenbach f.) L.O. Williams
TOOTHED HABENARIA
FL; MX, WtI, CtA
forma heatonii P.M. Brown- albino form
Brown, P.M. 2001. NA Native Orchid Journal 7(1): ined.
Habenaria quinqueseta (Michaux) Eaton
MICHAUX'S ORCHID
SC - FL w to TX
Habenaria repens Nuttall
WATER SPIDER ORCHID
NC - FL w to seAR, TX; MX, WtI, CtA
Harrisella porrecta (Reichenbach f.) Fawcett & Rendle
SYN: Campylocentrum porrectum Reichenbach f.
LEAFLESS HARRISELLA
FL; MX, CtA, WtI
Ackerman., J.D.. . 1995. Orchids of Puerto Rico and the Virgin Islands. p.
87..
Hexalectris grandiflora (A. Richard & Galeotti) L.O. Williams
GREENMAN'S CRESTED CORALROOT
TX; MX
Hexalectris nitida L.O. Williams
SHINING CRESTED CORALROOT
TX; MX
Hexalectris revoluta Correll
RECURVED CRESTED CORALROOT
AZ,TX; MX
Hexalectris spicata (Walter) Barnhardt var. spicata
CRESTED CORALROOT
AR, MO; sIL - MD s to FL - TX; MX
CHECKLIST
78
forma albolabia P.M. Brown - white flowered form
Brown, P.M.. 1995. NA Native Orchid Journal 1(1): 10.
Hexalectris spicata (Walter) Barnhardt var. arizonica (S. Watson) Catling &
Engel
ARIZONA CRESTED CORALROOT
AZ, TX; MX
Catling, P.M. & V.S. Engel. 1993. Lindleyana 8(3): 119-126.
Hexalectris warnockii Ames & Correll
TEXAS PURPLE-SPIKE
AZ; TX
Ionopsis utricularioides (Swartz) Lindley
DELICATE IONOPSIS
sFL; WtI, CtA, nSA
Isotria medeoloides (Pursh) Rafinesque
SMALL WHORLED POGONIA
MI - ME s to MO - GA
Isotria verticillata Rafinesque
LARGE WHORLED POGONIA
MI - ME s to TX - FL
Laelia rubescens Lindley
PALE LAELIA*
sFL
Lepanthopsis melanantha (Reichenbach f.) Ames
CRIMSON LEPANTHOPSIS
sFL; WtI
Lophiaris carthagenensis (Jacquin) G.A. Braem
SPREAD-EAGLE OORCHID
sFL; WtI, CtA, nSA
Braem, G.A. 1993. Schlechteriana 4:17
Lophiaris maculata (Aublet) Ackerman
CHECKLIST
79
SPOTTED MULE-EARED ORCHID
sFL; WtI, CtA, nSA
forma flavovirens (P.M. Brown) P.M. Brown - unspotted with a yellow-
green base
Ackerman, J. 2000. Lindleyana 15(2): 92-93.
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 132.
2000. NA Native Orchid Journal 6(4):
Liparis elata Lindley
SYN: Liparis nervosa (Thunberg) Lindley in part
TALL TWAYBLADE
FL; WtI, MX
Liparis liliifolia (Linnaeus) Richard
LILY-LEAVED TWAYBLADE
MN - VT s to AR - GA
forma viridiflora Wadmond - green-flowered form
Liparis loeselii (Linnaeus) Richard
LOESEL'S TWAYBLADE; FEN ORCHIS
sWA, MT; MAN - NS s to MS, sAM; Eurasia
HYBRID:
Liparis xjonesii S. Bentley
JONES' HYBRID TWAYBLADE
Bentley, S. 2000. Native Orchids of the Southern Appalachians, pp. 138-
139.
Listera auriculata Wiegand
AURICLED TWAYBLADE
ON - NF s to MI - ME
forma trifolia (Lepage) Lepage - three-leaved form
Listera australis Lindley
SOUTHERN TWAYBLADE
PQ, NS, NB - FL w to TX
forma scottii P.M. Brown - many leaved form
trifolia P.M. Brown - three-leaved form
CHECKLIST
80
viridis P.M. Brown - green-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 11.
2000. NA Native Orchid Journal 6(1):
Listera borealis Morong
NORTHERN TWAYBLADE
AK - NF s to UT, CO
forma trifolia Lepage - three-leaved form
Listera caurina Piper
NORTHWESTERN TWAYBLADE
sAK - ALB s to CA, WY
Listera convallarioides (Swartz) Nuttall
BROAD-LIPPED TWAYBLADE
swAK; BC - NF s to CA, WY e to nMI - nME
forma trifolia P. M. Brown - three-leaved form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 11
Listera cordata (Linnaeus) R. Brown var. cordata
HEART-LEAVED TWAYBLADE
AK - NF s to CA, NM in RM; NC in AM; Eurasia
forma disjuncta Lepage - alternate-leaved form
trifolia P. M. Brown - three-leaved form
variegata P. M. Brown - variegated-leaved form
viridens P.M. Brown - green-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 11; 1(4): 288.
Listera cordata (Linnaeus) R. Brown var. nephrophylla (Rydberg) Hulten
WESTERN HEART-LEAVED TWAYBLADE
RM w to CA n to AK
forma rubescens P.M. Brown - reddish-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 240; 1(4): 288.
Listera ovata (Linnaeus) Aiton f.
COMMON TWAYBLADE*
ON
Listera smallii Wiegand
SMALL'S TWAYBLADE
nNJ; PA - GA
forma variegata P.M. Brown - variegated-leaved form
Brown, P.M. 1995. NA Native Orchid Journal 1(4): 289.
CHECKLIST
81
Hybrid:
Listera xveltmanii Case
VELTMAN'S TWAYBLADE
(L. auriculata x L. convallarioides)
Macradenia lutescens R. Brown
TRINIDAD MACRADENIA
sFL; WtI, nSA
Malaxis bayardii Fernald
BAYARD'S ADDER'S-MOUTH
NS; MA - NC w to OH
Catling, P.M. 1991. Lindleyana 6(1): 3-23.
Luer, C.A. 1975. Nat. Orchids US & Can. Pl. 79:5.
Malaxis brachypoda (Gray) Fernald
SYN: Malxis monophyllos (Linnaeus) Swartz. var. brachypoda (A. Gray)
Morris & Eames
WHITE ADDER'S-MOUTH
sAK - NF s to CA, CO, IN, PA
forma bifolia (Mousley) Fernald - two-leaved form
Malaxis corymbosa (S. Watson) Kuntze
CLUSTERED ADDER'S-MOUTH
AZ; MX
Malaxis diphyllos Chamisso
SYN: Malaxis monophyllos (Linnaeus) Swartz. var. diphyllos (Chamisso)
Luer
TWO-LEAVED ADDER'S-MOUTH
swAK
Malaxis paludosa (Linnaeus) Swartz
SYN: Hammarbya paludosa (Linnaeus) Kuntze
BOG ADDER'S-MOUTH
AK- wONT s to MN; nEurasia
Reeves, T. & L. M. Reeves. 1984. AOS Bulletin 53(12): 1280-1292.
CHECKLIST
82
Reeves, T. & L. M. Reeves. 1985. Rhodora. 87: 133-136.
Malaxis porphyrea (Ridley) Kuntze
PURPLE ADDER’S-MOUTH
AZ,NM; MX
in Luer, 1975 as Malaxis ehrenbergii (Reichenbach f.) Kuntze, p. 296,
pl. 81.
Todsen, T. SIDA 1997. 17: 637-638.
Malaxis soulei L.O. Williams
SYN: Malaxis macrostachys (Lexara) Kuntze
RAT-TAILED ADDER'S-MOUTH
AZ, NM, TX; MX
Malaxis spicata Swartz
FLORIDA ADDER'S-MOUTH
VA - FL; BA; WtI
Malaxis tenuis (S. Watson) Ames
THIN ADDER'S-MOUTH
AZ, NM; MX
Malaxis unifolia Michaux
GREEN ADDER'S-MOUTH
MAN - NF s to TX - FL; MX
forma bifolia (Mousley) Fernald - two-leaved form
Malaxis wendtii Salazar
WENDT'S ADDER'S-MOUTH
TX; MX
Salazar, G. 1993. Orquidea(Mex.) 13(1-2): 281-284.
Todsen, T. SIDA 1995. 16(3): 591.
Todsen, T. SIDA 1997. 17: 637-638.
Maxillaria crassifolia (Lindley) Reichenbach f.
FALSE BUTTERFLY ORCHID
sFL; WtI, CtA, nSA
Maxillaria parviflora (Poeppig & Endlicher) Garay
SYN: Maxillaria conferta (Grisebach) C. Schweinfurth ex Leon
DENSELY-FLOWERED MAXILLARIA
CHECKLIST
83
sFL; WtI, MX, CtA, SA
Attwood, J. T. Lindleyana 8(1): 25-31.
Hammer, R. 1981. Fairchild Trop. Gard. Bulletin 36(3): 16-18.
McCartney, C. 1993. Florida Orchidist. 36(3): 25-29.
Mesadenus lucayanus (Britton) Schlechter
(Syn: Ibidium lucayanum Britton; Spiranthes lucayana (Britton) Cogniaux
COPPER LADIES'-TRESSES
FL; MX, BA, WtI, CtA
Brown, P.M. 200. NA Native Orchid Journal 6(4): 335-336.
Oeceoclades maculata (Lindley) Lindley
SPOTTED AFRICAN ORCHIS*
sFL; WtI, SA; Africa
Johnson, S.R. 1993. Lindleyana 8(2): 69-72.
Stern, W.S. 1988. AOS Bulletin 57(9): 960-971.
Oncidium floridanum Ames
SYN: Oncidium ensatum Lindley in part
FLORIDA ONCIDIUM
sFL; BA
see Lophiaris for additional species
Pelexia adnata (Swartz) Sprengl
SYN: Spiranthes adnata (Swartz) Bentham ex Fawcett
GLANDULAR LADIES'-TRESSES
sFL; WtI, CtA, MX, nSA
Hammer, R. L. 1981. Fairchild Trop. Garden Bulletin 36(3): 16-18.
McCartney, C.L., Jr. 1983. Florida Orchidist 26(3): 124-129.
Piperia candida Morgan & Ackerman
SLENDER WHITE PIPERIA
sAK - CA
Morgan, R. & J. Ackerman. 1990. Lindleyana 5(4): 205-211.
Piperia colemanii Morgan & Glicenstein
COLEMAN'S PIPERIA
CA
Morgan, R. & L. Glicenstein. 1993. Lindleyana 8(2): 89-95.
Piperia cooperi (S. Watson) Rydberg
COOPER'S STOUT-SPIRE ORCHID
CHECKLIST
84
CA
Coleman, R. A. 1992. AOS Bulletin 61(2): 130-135.
Coleman, R.A. 1995. NA Native Orchid Journal 1(1): 75-78.
Piperia elegans (Lindley) Rydberg subsp. elegans
SYN: Piperia maritima Rydberg
ELEGANT PIPERIA
BC - CA; nID, wMT
Piperia elegans subsp. decurtata Morgan & Glicenstein
PT. REYES PIPERIA
CA
Morgan, R & L. Glicenstein. 1993. Lindleyana 8(2): 89-95.
Piperia elongata Rydberg
SYN: Piperia elegans var. elata (Jepson) Luer
LONG-SPURRED PIPERIA
BC - CA; nID, wMT
Piperia leptopetala Rydberg
LACE ORCHID
WA - CA
Piperia michaelii (E. Greene) Rydberg
SYN: Piperia elongata var. michaelii (Greene) Ackerman
MICHAEL'S PIPERIA
OR - CA
Piperia transversa Suksdorf
FLAT-SPURRED PIPERIA
BC - CA
Piperia unalascensis (Sprengel) Rydberg
ALASKA PIPERIA
AK - CA w. to MT; nNM ; ON, MI, PQ
Piperia yadonii R. Morgan & J. Ackerman
YADON'S PIPERIA
CA
Morgan, R & J. Ackerman. 1990. Lindleyana 5(4): 205-211.
Platanthera aquilonis Sheviak
CHECKLIST
85
SYN: Platanthera hyperborea (Linnaeus) Lindley in part
NORTHERN GREEN BOG ORCHIS
AK - NF s to CA, NM, IA e to MA
forma alba (Light) P.M. Brown - albino form
Brown, P.M. 2000. NA Native Orchid Journal 6(1): 43.
Light, M.S. & M. MacConaill. 1989. Lindleyana 4(3): 158-160.
Sheviak, C.J. 1999. Lindleyana 14(4): 193-203
Platanthera blephariglottis (Willdenow) Lindley var. blephariglottis
NORTHERN WHITE FRINGED ORCHIS
MI - NF s to IL; OH - NJ
forma holopetala (Lindley) P.M. Brown - entire-lip form
Brown, P.M. 1988. Wild Flower Notes 3(1): 20.
Platanthera blephariglottis (Willdenow) Lindley var. conspicua (Nash) Luer
SOUTHERN WHITE FRINGED ORCHIS
se MA - FL w to TX
Platanthera brevifolia (Greene) Kranzlein
SYN: Platanthera sparsiflora (S. Watson) Schlecter var. brevifolia (Greene)
Luer
SHORT-LEAVED REIN ORCHIS
NM; MX
Platanthera chapmanii (Small) Luer emend. Folsom
CHAPMAN'S FRINGED ORCHIS
seGA, nFL, eTX
Folsom, J.P. 1984. Orquidea (Mex) .9(2): 344.
Platanthera chorisiana (Chamisso) Reichenbach f.
CHAMISSO'S ORCHID
AK - WA; Asia
Platanthera ciliaris (Linnaeus) Lindley
ORANGE FRINGED ORCHIS
MI - MA s to TX - FL
Platanthera clavellata (Michaux) Luer var. clavellata
LITTLE CLUB-SPUR ORCHIS
WI - ME s to TX - GA
forma slaughteri P.M. Brown - white-flowered form
Brown, P.M. 1995. NA Native Orchid Journal 1(3): 200.
Platanthera clavellata (Michaux) Luer var. ophioglossoides (Fernald) P.M.
Brown
CHECKLIST
86
NORTHERN CLUB-SPUR ORCHIS
ON - NF s to MI - MA
Brown, P.M. 1988. Wild Flower Notes 3(1): 21.
Platanthera cristata (Michaux) Lindley
ORANGE CRESTED ORCHIS
MA - FL w to TX
forma straminea P.M. Brown - pale yellow form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 12.
Platanthera dilatata (Pursh) Lindley var. dilatata
TALL WHITE NORTHERN BOG ORCHIS
AK - NF so to CA - NM; MN -IN, PA
Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour
BOG CANDLES
AK - WA e to UT, CO
Platanthera dilatata (Pursh) Lindley var. leucostachys (Lindley) Luer
SIERRA REIN-ORCHID
sAK - CA e to WY - nAZ
Platanthera flava (Linnaeus) Lindley var. flava
SOUTHERN TUBERCLED ORCHIS
swNS; MO - MD s to TX - FL
Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer
NORTHERN TUBERCLED ORCHIS
MN - NS s to MO - GA; s in AM
forma lutea (Boivin) Whiting & Catling - yellow-flowered form
Platanthera grandiflora (Bigelow) Lindley
LARGE PURPLE FRINGED ORCHIS
MN - NF s to WV - NJ; s in AM to GA
forma albiflora (Rand & Redfield) Catling - white-flowered form
bicolor P.M. Brown - bicolor-flowered form
carnea P.M. Brown - pink-flowered form
mentotonsa (Fernald) P.M. Brown - entire-lip form
Brown, P.M. 1988. Wild Flower Notes 3(1): 22.
Brown, .M. 1995. NA Native Orchid Journal 1(1): 12.
Stoutamire, W.P. 1974. Brittonia 26: 42-58.
Platanthera hookeri (Torrey) Lindley
HOOKER'S ORCHIS
forma abbreviata (Fernald) P.M. Brown - dwarfed form
MAN - NF s to IA - NJ
CHECKLIST
87
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 14.
Platanthera huronensis (Nuttall) Lindley
SYN: Platanthera hyperborea (Linnaeus) Lindley var. huronensis (Nuttall)
Luer
GREEN BOG ORCHIS
AK - NF s to CA - PA
Platanthera hyperborea (Linnaeus) Lindley var. gracilis (Lindley) Luer
LAXLY FLOWERED BOG ORCHIS
sAK - MT s to CA - CO
Platanthera hyperborea (Linnaeus) Lindley var. viridiflora (Chamisso) Luer
SYN: Platanthera convallariifolia (Fischer) Lindley
TALL ALASKA GREEN ORCHIS
AK
Platanthera integra (Nuttall) Lindley
YELLOW FRINGELESS ORCHIS
NJ - FL w to TX
Platanthera integrilabia (Correll) Luer
MONKEY-FACE
KY - NC s to MS - GA
Zettler, L.W. 1994. AOS Bulletin 63(6): 686-688
Zettler, L.W. & J.E. Fairey, III. 1992. Lindleyana 5(4): 212-217.
Platanthera lacera (Michaux) G. Don
GREEN FRINGED ORCHIS; RAGGED ORCHIS
MAN - NF s to OK - GA
Catling, P.M. 1997. Lindleyana. 12(2):79-88..
Platanthera leucophaea (Nuttall) Lindley
EASTERN PRAIRIE FRINGED ORCHIS
NE - ME s to OK - VA
Sheviak, C. J. & M. Bowles. 1986. Rhodora. 88: 267-290.
Platanthera limosa Lindley
THURBER'S BOG ORCHID
AZ-NM; MX
Platanthera macrophylla (Goldie) P.M. Brown
GOLDIE'S PAD-LEAVED ORCHIS
CHECKLIST
88
ON - NF s to MI - PA
Brown, P.M. 1988. Wild Flower Notes 3(1): 23.
Reddoch, A.H. & J. M. Reddoch 1993. Lindleyana. 8(4): 171-188.
Platanthera nivea (Nuttall) Luer
SNOWY ORCHIS
sNJ - FL w to TX
Platanthera obtusata (Banks ex Pursh) Lindley
BLUNT-LEAVED REIN ORCHIS
AK - NF s to CO, GtL, MA
forma collectanea (Fernald) P.M. Brown - dwarfed form
foliosa P.M. Brown - multiple-leaved form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 13-14.
Platanthera orbiculata (Pursh) Lindley
PAD-LEAVED ORCHIS
seAK, BC - NF s to WA - MD, s in AM to NC
forma lehorsii (Fernald) P.M. Brown - dwarfed form
trifolia (Mousley) P.M. Brown - three leaved form
Brown, P.M. 1995. NA Native Orchid Journal 1(1): 15.
Reddoch, A.H. & J. M. Reddoch. 1993. Lindleyana. 8(4): 171-188.
Platanthera pallida P.M. Brown
PALE FRINGED ORCHIS
eNY
Brown, P.M. 1993. Novon. 2(4): 308-311.
Platanthera peramoena (A. Gray) A. Gray
PURPLE FRINGLESS ORCHIS
MO - NJ s to MS - GA
Spooner, D.M. & J.S. Shelly. 1983. Rhodora. 85: 55-64.
Platanthera praeclara Sheviak & Bowles
WESTERN PRAIRIE FRINGED ORCHIS
WY - MN s to OK - MO
Sheviak, C. J. & M. Bowles. 1986. Rhodora. 88: 267-290.
Smith, W.R. 1995. Orchids of Minn. p. 136-137.
Platanthera psycodes (Linnaeus) Lindley
SMALL PURPLE FRINGED ORCHIS
MN - NF s to OH - NJ, s in AM to GA
forma albiflora (R. Hoffman) Whiting & Catling -
CHECKLIST
89
white-flowered form
ecalcarata (Bryan) P.M. Brown - spurless form
rosea P.M. Brown - pink -colored form
varians (Bryan) P.M. Brown - entire-lip form
Brown, P.M. 1988. Wild Flower Notes 3(1): 24.
Brown, P.M. 1995. NA Native Orchid Journal 1(4): 289.
Stoutamire, W.P. 1974. Brittonia .26: 42-58.
Platanthera purpurascens (Rydberg) Sheviak & Jennings
SHORT-SPURRED BOG ORCHIS
CA - NM n to CO
Sheviak & Jennings. 1997. NA Native Orchid Journal 3(4): 444-449.
Platanthera sparsiflora (S. Watson) Schlecter var. sparsiflora
FEW-FLOWERED REIN-ORCHIS
WA - CA; UT-NM; MX
Platanthera sparsiflora (S. Watson) Schlecter var. ensifolia (Rydberg) Luer
NARROW-LEAVED REIN-ORCHIS
WA-CA
Platanthera stricta Lindley
SYN: Platanthera saccata (Greene) Hulten
SLENDER BOG ORCHIS
AK - ALB s to CA - WY
Platanthera tipuloides (Linnaeus). Lindley var. behringiana (Rydberg)
Hulten
BEHRING ORCHID
AK; neAsia
Platanthera zothecina (Higgins & Welsh) Kartesz & Ghandi
CLOISTERED BOG ORCHID
UT, CO, AZ
Higgins, L.C. & S. L. Welsh. 1986. Great Basin Naturalist. 46: 259.
in Long, J.C. 1970. Nat. Orchids Col. as Habenaria sparsiflora S.
Watson, p. 22.
Hybrids:
Platanthera xandrewsii (Niles) Luer
SYN: Platanthera lacera var. terrae-novae (Fernald) Luer
ANDREWS' FRINGED ORCHIS
(P. lacera x P. psycodes)
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.
CHECKLIST
90
Platanthera xbicolor (Rafinesque) Luer
BICOLOR FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. ciliaris)
Platanthera xcanbyi (Ames) Luer
CANBY'S FRINGED ORCHIS
(P. blephariglottis var. conspicua x P. cristata)
Platanthera xchannellii Folsom
CHANNELL'S FRINGED ORCHIS
(P. ciliaris x P. cristata)
Folsom, J.P. 1984. Orquidea (Mex) 9(2): 344.
Platanthera xcorrellii Schrenck
CORRELL'S REIN ORCHIS
(P. hyperborea x P. stricta)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263.
Schrenck, W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xestesii Schrenck
ESTES REIN ORCHIS
(P. dilatata var. albiflora x P. stricta)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263.
Schrenck, W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xkeenanii P.M. Brown
KEENAN'S FRINGED ORCHIS
(P. grandiflora x P. lacera)
Brown, P.M. 1993. A Field Guide to the Orchids of N.E. & N.Y. p. 189.
Catling, P.M. & V. Catling. 1994. Lindleyana 9(1): 19-32.
Platanthera xlassenii Schrenk
LASSEN REIN ORCHIS
(P. leucostachys x P. sparsiflora)
Schrenck, W.J. 1975. Die Orchidee. 26: 258-263.
Schrenck, W.J. 1978. AOS Bulletin. 47(5): 429-437.
Platanthera xmedia (Rydberg) Luer
INTERMEDIATE REIN ORCHIS
(P. hyperborea x P. dilatata)
Platanthera xreznicekii Catling, Brownell & Allen
CHECKLIST
91
REZNICEK'S ORCHID
(P. leucophaea x P. psycodes)
Catling, Brownell & Allen. 1999. Lindleyana 14(2):77-86.
Platanthera xvossii Case
VOSS' REIN ORCHIS
(P. blephariglottis var. blephariglottis x P. clavellata var.
ophioglossoides)
Case, F. W. 1983. Michigan Botanist. 22: 141-144.
Platytheles querciticola (Lindley)Garay
SYN: Erythrodes querciticola (Lindley) Ames
LOW GROUND ORCHID
TX - FL
Platytheles sagreana (A. Richard) Garay
CUBAN GROUND ORCHID
sFL; WtI
Brown, P.M. 1998. NA Native Orchid Journal 5(1):3
Pleurothallis gelida Lindley
FROSTED PLEUROTHALLIS
sFL; WtI, CtA, nSA
Pogonia ophioglossoides (Linnaeus) Ker-Gawler
ROSE POGONIA
ON - NF s to TX - FL
forma albiflora Rand & Redfield - white-flowered form
forma brachypogon (Fernald) P.M. Brown - short-bearded form
Brown, P.M. 1998. NA Native Orchid Journal 6(4): 339.
Polyradicion see Dendrophylax lindenii
Polystachya concreta (Jacquin) Garay & Sweet
SYN: Polystachya flavescens (Lindley) J.J. Small
PALE-FLOWERED POLYSTACHYA
sFL; WtI, CtA, n + cSA
Ponthieva brittoniae Ames
SYN: Ponthieva racemosa (Walter) C. Mohr var. brittonae (Ames) Luer
MRS. BRITTON'S SHADOW-WITCH
sFL; BA
McCartney, C.L., Jr. 1995. NA Native Orchid Journal 1(2): 106-116.
CHECKLIST
92
Ponthieva racemosa (Walter) Mohr
SHADOW-WITCH
VA - FL w to TX; MX, WtI, CtA, SA
Prescottia oligantha (Swartz) Lindley
SMALL PRESCOTTIA
sFL; WtI, CtA, n + cSA
Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small)
W.E. Higgins
SYN.:Encyclia boothiana (Lindley) Dressler var. erythronioides (Small)
Luer
FLORIDA DOLLAR ORCHID
sFL; CtA, WtI
Brown, P.M. 1998. NA Native Orchid Journal 4(1): 52.
Higgins, W.E. 1998. Phytologia 85(5):370-383.
Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E.
Higgins
SYN: Anacheilum cochleatum (Linnaeus) Small var. triandrum (Ames)
Saleuda et al.
Encyclia cochleata (Linnaeus) Dressler var. triandra (Ames) Dressler
FLORIDA CLAM-SHELL ORCHID
sFL; WtI, CtA, nSA
forma albidoflava (P.M.Brown) P. M. Brown - pale colored form
Brown, P.M. 1995. NA Native Orchid Journal 1(2): 131.
1998. NA Native Orchid Journal 4(1): 53.
Higgins, W.E. 1998. Phytologia 85(5):370-383.
Prosthechea pygmaea (Hooker) W.E. Higgins
SYN: Encyclia pygmaea (Hooker) Dressler
Hormidium pygmaceum (Hooker) Bentham ex Hemsley
DWARF BUTTERFLY ORCHID
sFL; WtI, CtA, nSA
Brown, P.M. 1998. NA Native Orchid Journal 4(1): 52.
Higgins, W.E. 1998. Phytologia 85(5):370-383.
Pseudorchis straminea (Fernald) Soo
SYN: Habenaria albida (Linnaeus) R. Brown var. straminea (Fernald)
Morris & Ames
Platanthera albida (Linnaeus) Lindley var. straminea (Fernald) Luer
CHECKLIST
93
Pseudorchis albida (Linnaeus) Love & Love subsp. straminea
(Fernald) Love & Love
NEWFOUNDLAND ORCHIS
NF; wPQ; Greenland, Iceland
Reinhammar, L. 1995. Nordic Journal of Botany 15(5): 469-481.
1997. NA Native Orchid Journal 3(4): 407-425.
Pteroglossaspis ecristata (Fernald) Rolfe
SYN: Eulophia ecristata (Fernald) Ames
CRESTLESS PLUME ORCHID
NC-FL w to LA; Cuba forma flava P.M. Brown - yellow-flowered form
Brown, P.M. 2000. NA Native Orchid Journal 6(1): 64.
Sacoila lanceolata (Aublet) Garay var. lanceolata
SYN: Spiranthes lanceolata (Aublet) Leon
Spiranthes orchioides (Swartz) A. Richard
Stenorrhynchos lanceolatum (Aublet) Richard ex Sprengel
LEAFLESS BEAKED ORCHID
FL; MX, BA, WtI, CtA, SA
forma albidaviridis Catling & Sheviak - white/green flowered form
folsomii P.M. Brown - golden-bronze form
FOLSOM'S GOLDEN LADIES'-TRESSES
Brown, P.M. 1999. NA Native Orchid Journal 5(2): 169-173.
Catling, P. M. & C. J. Sheviak. 1993. Lindleyana. 8(2): 77-81.
Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Saluda,
Wunderlein et Hansen
SYN: Spiranthes lanceolata (Aublet) Leon var. paludicola Luer
FAHKAHATCHEE BEAKED ORCHID
sFL
forma lutea P.M. Brown ined. - yellow flowered form
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana 8(2): 77-81.
Brown, P.M. 2001. NA Native Orchid Journal 7(1): ined.
Sacoila squamulosa (H.B.K.) Garay
HOARY LEAFLESS BEAKED ORCHID
Syn: Sacoila lanceolata (Aublet) Garay var. squamulosa (H.B.K.)
Szlachetko
Spiranthes squamulosa (H.B.K.) Leon
Stenorrhynchos squamulosum (H.B.K.) Sprengel
FL; W Indies
Brown, P.M. 2000. NA Native Orchid Journal 6(4):
CHECKLIST
94
Schiedeella arizonica P.M. Brown
RED-SPOT LADIES'-TRESSES
AZ - TX
in Luer, 1975 as Spiranthes parasitica (A. Richard & Galeotti) Schlecter;
p. 128, pl. 29:4-9.
Brown, P.M. 1996. NA Native Orchid Journal 2(1): 66-68. 2000. NA Native Orchid Journal 6(1): 3-17.
Spathoglottis plicata Blume
PURPLE JAVANESE ORCHID
ecFL
Spiranthes amesiana Eaton
AMES' LADIES'-TRESSES
FL; CtA
Spiranthes brevilabris Lindley
SHORT-LIPPED LADIES'-TRESSES; TEXAS LADIES'-TRESSES
TX - SC & FL
Spiranthes casei Catling & Cruise var. casei
CASE'S LADIES'-TRESSES
ON - NS s to MI, PA, NY - ME
in Luer, 1975 as Spiranthes intermedia Ames; p. 108, pl. 23:3-5.
Catling, P.M. & J.E. Cruise. 1974. Rhodora 76: 256-536.
Spiranthes casei Catling & Cruise var. novaescotiae Catling
CASE'S NOVA SCOTIAN LADIES'-TRESSES
NS
Catling, P.M. 1981. Can. J. Bot. 59: 1253-1270
Spiranthes cernua (Linnaeus) L.C. Richard
NODDING LADIES'-TRESSES
SD - NS s to TX - GA
Sheviak, C.J. 1991. Lindleyana 6(4): 228-234.
Spiranthes delitescens Sheviak
CIENEGAS LADIES'-TRESSES
AZ
in Luer, 1975. as Spiranthes graminea Lindley; p. 129, pl. 29:1-3
McClaren, M.C. 1996. NA Native Orchid Journal 2(2): 151-169.
McClaren, M.C. & P.C. Sundt.1992. Southwestern Naturalist 37: 299-333.
Sheviak, C.J. 1990. Rhodora 92: 213-231.
CHECKLIST
95
Spiranthes diluvialis Sheviak
UTE LADIES'-TRESSES
WA; UT, NV, CO, WY, MT, NE
Arf, A. M. 1994. Aquilegia 18(2): 1,4-5.
________ 1995. NA Native Orchid Journal 1(2): 117-128.
Sheviak, C.J. 1984. Brittonia 36: 8-14.
Spiranthes eatonii Ames ex P.M. Brown
EATON'S LADIES'TRESSES
NC - FL w to eLA
Brown, P.M. 1999. NA Native Orchid Journal 5(1):3
Spiranthes. floridana (Wherry) Cory
FLORIDA LADIES'-TRESSES
TX - NC
Spiranthes infernalis Sheviak
ASH MEADOWS LADIES'-TRESSES
NV
Sheviak, C.J. 1989. Rhodora 91: 225-234.
Spiranthes lacera Rafinesque var. lacera
NORTHERN SLENDER LADIES'-TRESSES
ALB - NS s to MO - VA
Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer
SYN: Spiranthes gracilis Bigelow
SOUTHERN SLENDER LADIES'-TRESSES
KS, MI - ME s to TX - GA
Spiranthes laciniata (Small) Ames
LACE-LIPPED LADIES'-TRESSES
sNJ - FL w to TX
Spiranthes longilabris Lindley
LONG-LIPPED LADIES'-TRESSES
VA - FL w to TX
Spiranthes lucida (H.H. Eaton) Ames
SHINING LADIES'-TRESSES
WI - NS s to KS - WV
Spiranthes magnicamporum Sheviak
GREAT PLAINS LADIES'-TRESSES
CHECKLIST
96
ND - sON s to NM, TX - GA; swVA
Spiranthes ochroleuca (Rydberg) Rydberg
YELLOW LADIES'-TRESSES
MI - NS s to KY - VA
Spiranthes odorata (Nuttall) Lindley
SYN: Spiranthes cernua var. odorata (Nuttall) Correll
FRAGRANT LADIES'-TRESSES
NJ - FL w to TX
Spiranthes ovalis Lindley var. ovalis
SOUTHERN OVAL LADIES'-TRESSES
AR - TX e to FL
Catling, P.M. 1983. Brittonia 35: 120-125.
Spiranthes ovalis Lindley var. erostellata Catling
NORTHERN OVAL LADIES'-TRESSES
WI - sON s to LA - FL
Catling, P.M. 1983. Brittonia 35: 120-125.
Spiranthes parksii Correll
NAVASOTA LADIES'-TRESSES
TX
Catling, P.M. & K. L. McIntosh. 1979. SIDA 8: 188-193.
Spiranthes porrifolia Lindley
WESTERN LADIES'-TRESSES
WA - CA e to NV
Spiranthes praecox (Walter) S. Watson
GIANT LADIES'-TRESSES
NY - FL w to TX
forma albolabia Brown & McCartney - white-lipped form
Brown,P.M. 1995. NA Native Orchid Journal 1(1): 13.
Spiranthes romanzoffiana Chamisso
HOODED LADIES'-TRESSES
AK - NF s to CA, nNM, IN, PA; Ireland
Spiranthes torta (Thunberg) Garay & Sweet
SYN: Spiranthes tortilis (Swartz) L.C. Richard
SOUTHERN LADIES'-TRESSES
sLA , sFL; BA, WtI, CtA
CHECKLIST
97
Spiranthes tuberosa Rafinesque
SYN: Spiranthes grayi Ames
LITTLE LADIES'-TRESSES
MI - MA s to TX - FL
Spiranthes vernalis Engler & Gray
GRASS-LEAVED LADIES'-TRESSES
NE - NH s to TX - FL; MX
Hybrids:
Spiranthes xborealis P.M. Brown
NORTHERN HYBRID LADIES'-TRESSES
(S. casei var. casei x S. ochroleuca)
Brown, P.M. 1995. N.A Native Orchid Journal 1(4): 290. Spiranthes xfolsomii P.M. Brown (S. longilabris x S. odorata) FOLSOM"S HYBRID LADIES"TRESSES Brown, P.M. 2000. NA Native Orchid Journal 6(1): 16.
Spiranthes xitchetuckneensis P.M. Brown
ITCHETUCKNEE SPRINGS LADIES'-TRESSES
(S. ovalis var. ovalis x S. odorata)
Brown, P.M. 1999. N.A Native Orchid Journal 5(4): 358-367.
Spiranthes xintermedia Ames
HYBRID LADIES'-TRESSES
(S. lacera var. gracilis x S. vernalis)
Catling, P.M. 1978. Rhodora 80: 377-389.
Spiranthes xmeridionalis P.M. Brown
SOUTHERN HYBRID LADIES'-TRESSES
(S. vernalis x S. praecox)
Brown, P.M. 1999. N.A Native Orchid Journal 5(4): 358-367.
2000. N.A Native Orchid Journal 6(2): 139.
Spiranthes xsimpsonii Catling & Sheviak
SIMPSON'S LADIES'-TRESSES
(S. lacera var. lacera x S. romanzoffiana)
Catling, P.M. & C.J. Sheviak. 1993. Lindleyana. 8(2): 78-80.
CHECKLIST
98
Stenorrhynchos michuacanum (Llave & Lexara) Lindley
SYN: Spiranthes michuacana (Lexara) Hemsley
MICHOACAN LADIES'-TRESSES
AZ, TX; MX
Coleman, R.A. 1996. Orchids 65(12): 1284-1287.
Tipularia discolor (Pursh) Nuttall
CRANE-FLY ORCHIS
MI - MA s to TX - FL
forma viridifolia P.M. Brown
Brown, P.M. 2000. NA Native Orchid Journal 6(4):336-337.
Tolumnia variegata (Swartz) Braem
SYN: Oncidium varigatum (Swartz)
Oncidium bahamense Nash ex Britton & Millspaugh
Tolumnia bahamensis (Nash ex Britton & Millspaugh) G.J. Braem
FLORIDA VARIEGATED ONCIDIUM
se cFL, BA
Ackerman, J. 2000. Lindleyana 15(2): 93.
Sauleda, R.P. & R.M. Adams. 1989. Rhodora 91(866): 188-200.
Triphora amazonica Schlechter.
SYN: Triphora latifolia Luer f
WIDE-LEAVED TRIPHORA
FL; WtI
Ackerman, J. 2000. Lindleyana 15(2): 93-94.
Triphora craigheadii Luer
CRAIGHEAD'S TRIPHORA
FL
Triphora gentianoides (Swartz) Ames & Schlecter
LEAST FLOWERED TRIPHORA
seFL; WtI, CtA, nSA
Triphora trianthophora (Swartz) Rydberg subsp. trianthophora
THREE BIRD'S ORCHIS; NODDING POGONIA
[including var. schaffneri Campbell]
IA - ME s to TX - FL
forma albidoflava Keenan - white-flowered form
caerulea P.M. Brown ined.- blue-flowered form
rossii P.M. Brown - multi-color form
Brown, P.M. 1999. NA Native Orchid Journal 5(1): 5
CHECKLIST
99
2001. NA Native Orchid Journal 7(1): ined.
Keenan, P. 1992. Rhodora 94: 38-39.
Medley, M.E. 1991. Selbyana 12: 102-103.
Triphora rickettii Luer
RICKETT'S TRIPHORA
FL
Tropidia polystachya (Swartz) Ames
MANY-FLOWERED TROPIDIA
sFL; WtI, CtA, nSA
Vanilla barbellata Reichenbach f.
WORM-VINE; LEAFLESS VANILLA
sFL; WtI
Vanilla dilloniana Correll
DILLON'S VANILLA
sFL; WtI
Vanilla mexicana Miller
SYN: Vanilla inodora Schiede
SCENTLESS VANILLA
sFL; WtI, CtA, nSA
Vanilla phaeantha Reichenbach f.
OBLONG-LEAVED VANILLA
sFL; WtI
Vanilla planifolia Jackson in Andrews
COMMERCIAL VANILLA
sFL; WtI, CtA, n + wSA
Vanilla pompona Schiede
SOUTHERN VANILLA*
sFL
Zeuxine strateumatica (Linnaeus) Schlechter
LAWN ORCHID*
GA - FL; PR; se Asia
ANNOTATIONS TO THE CHECKLIST
100
ANNOTATIONS TO THE CHECKLIST
Paul Martin Brown
The following annotations are designed to help the reader with recently revised taxa, call attention to significant range extensions and clarify some finer points of taxonomic controversy. Full reference citations are included within the Checklist. Several persons contributed to the information in the annotations including (but not limited to) Chuck Sheviak, Paul Catling, Dick Wunderlin, Bruce Hansen, Bob Dressler, Chuck McCartney and John Beckner. Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis (Lindley) Ames Corallorhiza odontorhiza var. pringlei (Greenman) Freudenstein Freudenstein's recent mongraph on Corallorhiza details the differences and ranges. 1997.
ANNOTATIONS TO THE CHECKLIST
101
Cyclopogon cranichoides (Grisebach) Schlechter Cyclopogon elatus (Swartz) Schlechter At one time viewed as Spiranthes, more recently as Beadlea and, currently by Ackerman (1995) as Cyclopogon; based on the presence of intermediate taxa between Beadlea and Cyclopogon, with the latter name having priority Cypripedium kentuckiense C. F. Reed PURLOINED LADY'S-SLIPPER; IVORY-LIPPED LADY'S-SLIPPER Well known, but recently described species with a long (and convoluted) taxonomic history. See Brown, 1995 for details. Sheviak suggests purloined lady's-slipper for the common name in reference to the original discovery and subsequent publication! Recent range extension to eastern Virginia Cypripedium parviflorum Salisbury var. parviflorum Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak Cypripedium parviflorum Salisbury var. pubescens Again a long, and often confusing, taxonomic history, but Sheviak (1996) has apparently sorted this out with a great degree of satisfaction. Cyrtopodium paranaense Schlecter This is the only correct identification of the 'other Cytopodium' in Florida See Roger Hammer's excellent and detailed article of misidentification and eventual determination (1997). Cyrtopodium andersonii does not occur - either native or introduced - in Florida.
ANNOTATIONS TO THE CHECKLIST
102
Dactylorhiza praetermissa (Druce) Soo One of two taxa of dactylorchids present in Newfoundland. Some taxonomic confusion, but recent European work uses this epithet rather than D. majalis var. praetermissa. D. praetermissa var. junialis Recent discovery in eastern Newfoundland. Meades, 1995; Clase & Meades, 1996. Encyclia rufa Based upon a collection of Small from Brevard Co., Florida. Epidendrum floridense Hagsater After years of confusion with the genus Hagsater (1993) has sorted them all out and described a number of new taxa, most of them geographically isolated. Epidendrum floridense is endemic to Florida and all specimens of E. difforme are E. floridense. Govenia sp. See Ed Greenwood's work in for details of why this is not G. utriculata and perhaps cannot be accurately identified. 1991/1996.
ANNOTATIONS TO THE CHECKLIST
103
Habenaria odontopetala Reichenbach f. SYN: Habenaria strictissima Reichenbach f. var. odontopetala (Reichenbach f.) L.O. Williams Habenaria floribunda Lindley Although some Florida botanists are using Habenaria floribunda based on the possibility that there may be several taxa (3 in Florida) present within what most are calling H. odontopetala. If this is correct and there are also additional taxa in the Caribbean and Central and South America, until all of these can be sorted out they would fall under the oldest name for the group, which is H. floribunda. Much work is needed to resolve this situation. Liparis nervosa (Thunberg) Lindley SYN: Liparis elata Lindley in part A simple case of lumping and splitting. The broader taxon is L. nervosa. If one recognizes New World plants as separate from Old World plants, L. elata is applicable. Garay (1971) concluded that they are synonymous. Malaxis porphyrea (Ridley) Kuntze Malaxis wendtii Salazar Malaxis ehrenbergii (Reichenbach f.) Kuntze A tale of three red-flowered adder's-mouths from the Southwest U.S. A bit confusing to follow because of several publications and subsequent reassessments. Here is the recent chronology:
red-flowered Malaxis from Arizona, New Mexico, Texas and Mexico all known as Malaxis ehrenbergii
Salazar (1993) publishes Malaxis wendtii to separate two taxa in Mexico.
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Todsen (1995) reassesses the US plants and publishes note that they should be referred to M. wendtii as are those in northern Mexico.
Closer examination by Coleman and Todsen of Arizona and New Mexico material show they are not M. wendtii nor are they the more southerly M. ehrenbergii. Subsequently they are identified as M. porphyrea, with the type specimen from Arizona. These include plants from Arizona and New Mexico. 1997.
Todsen determines that plants from the Big Bend area of west Texas are actually M. wendtii. 1997.
Malaxis soulei L.O. Williams Prior and valid name for M. macrostachya Platanthera pallida P.M. Brown Recently describe from eastern Long Island and part of the P. cristata/blephariglottis/ciliaris complex. Plants reduced to synonymy with P. cristata and at one time with P. xcanbyi (hybrid between P. cristata and P. blephariglottis). Recent unpublished work form the University of Wisconsin shows that they have parentage with P. cristata and P. blephariglottis, but not in the same proportions as P. xcanbyi. P. pallida forms large, reproducing colonies whereas P. xcanbyi occurs as scattered individuals with both parents. A similar situation can be found in P. chapmanii and P. xchannellii. 1993. Platanthera purpurascens (Rydberg) Sheviak & Jennings
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See Sheviak and Jennings (1997) recent article on this new combination Prosthechea boothiana (Lindley) W.E. Higgins var. erythronioides (Small) W.E. Higgins Prosthechea cochleata (Linnaeus) W.E. Higgins var. triandra (Ames) W.E. Higgins Prosthechea pygmaea (Hooker) W.E. Higgins All formerly in the genus Encyclia. From Wes Higgins at the University of Florida: An ongoing systematic study of the genus Encyclia based on holomorphology has determined that the genus is neither morphologically cohesive nor monophyletic. In a preliminary molecular study, analysis of the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA supports the morphological conclusion that the Encyclia subgenus Osmophytum clade should be raised to the generic level because these species are sister to the Cattleya-Laelia clade not to Encyclia subgenus Encyclia. However, the monophyly of the three currently recognized subgenera of Encyclia i.e., Encyclia subg. Osmophytum, Encyclia subg. Encyclia, and Encyclia subg. Dinema, is supported by cladistic analysis of both morphological and molecular data. Encyclia subgenus Osmophytum
is raised to generic level and treated as Prosthechea. 1998. Pseudorchis straminea (Fernald) Soo Recent analysis by Lars-Gunner Reinhammar (1995/1997) indicates that two distinct species should be recognized. Ours in North America is the above. Schiedeella fauci-sanguinea (Dod) Burns-Balogh Although plants of Schiedeella parasitica in the southwestern US may be referable to this taxon, research is underway that indicates that it most likely is neither S.
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fauci-sanguinea nor S. parasitica and may be in need of a new description. 1996. Spiranthes sinensis (Persoon) Ames Recent and most curious addition to the orchid flora of North America. See John Beckner's detailed account. 1996. Spiranthes amesiana Schlechter Included by Wunderlin et al. in the Florida Atlas based upon the single specimen at AMES. Included in synonymy with S. torta by most other authorities. Whether it was a single anomaly or has simply been overlooked remains to be determined. Fieldwork in south Florida should help to solve this mystery. 1996. Spiranthes brevilabris Lindley Spiranthes floridana (Wherry) Cory Recognized by Wunderlin et al. as separate species. If based purely on degree of pubescence and range, that hardly seems sufficient. If there are other characters that separate the taxa, then perhaps they can be viewed as two species. Has been suggested that perhaps S. brevilabris is a hybrid of S. vernalis and S. floridana. 1996.
RECENTLY DESCRIBED OR TRANSFERRED TAXA
In the past 25 years many new species have been
described for North America, as well as several varieties elevated to species level. The following lists them with
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their date of publication. This list does not include forma.
New Species Calopogon oklahomensis 1995 Cypripedium kentuckiense 1981 Malaxis wendtii 1993 Piperia candida 1990 Piperia colemanii 1993 Piperia elegans subsp. decurtata 1993 Piperia yadonii 1990 Platanthera pallida 1993 Platanthera praeclara 1986 Platanthera zothecina 1986 Spiranthes casei var. casei 1974 Spiranthes casei var. novaescotiae 1981 Spiranthes ovalis var. erostellata 1983 Spiranthes delitescens 1990 Spiranthes diluvialis 1984 Spiranthes inferrnalis 1989 New status Cypripedium parviflorum var. makasin Corallorhiza odontorhiza var. pringlei Hexalectris spicata var. arizonica Restored to the species level, from either varietal status or synonymy Cleistes bifaria Cypripedium yatabeanum Malaxis bayardii Malaxis diphyllos
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Malaxis porphyrea Malaxis soulei Platanthera macrophylla Platanthera purpurascens Ponthieva brittoniae Pseudorchis straminea Spiranthes amesiana Spiranthes floridana Spiranthes odorata Spiranthes ochroleuca From the hybrid level Platanthera chapmanii New Hybrids Cypripedium xalaskanum Platanthera xchannellii Platanthera xkeenanii Platanthera xvossii Spiranthes xborealis Spiranthes xsimpsonii Excluded species: In the Spring 1997 issue of The Palmetto Chuck McCartney deals will a number of Florida species which have had dubious documentation in the United States. Those species are excluded from the checklist. Brassavola nodosa Leochilus labiatus Maxillaria sanguinea Restrepiella ophiocephala Tetramicra caniculata
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Those species which are considered introduced or adventive: This often becomes a difficult assessment. Some species are well documented, such as an escaped Bletilla or Cyrtopodium, others are more difficult to assess. There could perhaps be additional species listed here, such as Pelexia adnata, if we consider that they simply may have 'blown over' from the Caribbean. See note on Spiranthes sinensis in the next article. Bletilla striata Cyrtopodium paranaense Dactylorhiza cf. fuchsii Epidendrum radicans Epipactis atrorubens Epipactis helleborine Gymnadenia conopsea Listera ovata Oeceoclades maculata Zeuxine strateumatica Those species formerly included in the genus Habenaria and most often found within that species in the majority of the literature. It is implied that all subspecies and varieties are included within the species, and are listed only if the name is significantly different. This list is not meant to be exhaustive, but contains the vast majority of synonyms used with Habenaria in the bulk of the literature. Habenaria albida var. straminea = Pseudorchis straminea Habenaria blephariglottis = Platanthera blephariglottis
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Habenaria brevifolia = Platanthera brevifolia Habenaria chorisiana = Platanthera chorisiana Habenaria ciliaris = Platanthera ciliaris Habenaria clavellata = Platanthera clavellata Habenaria cooperi = Piperia cooperi Habenaria correlliana = Platanthera integrilabia Habenaria cristata = Platanthera cristata Habenaria dilatata = Platanthera dilatata Habenaria elegans = Piperia elegans Habenaria elegans var. elata = Piperia elongata Habenaria elongata = Piperia elongata Habenaria flava = Platanthera flava Habenaria grandiflora = Platanthera grandiflora Habenaria hookeri = Platanthera hookeri Habenaria huronensis = Platanthera huronensis Habenaria integra = Platanthera integra Habenaria integrilabia = Platanthera integrilabia Habenaria lacera = Platanthera lacera Habenaria leucophaea = Platanthera leucophaea Habenaria leucophaea var. praeclara = Platanthera praeclara Habenaria limosa = Platanthera limosa Habenaria macrophylla = Platanthera macrophylla Habenaria maritima = Piperia elegans Habenaria michaelii = Piperia michaelii Habenaria nivea = Platanthera nivea Habenaria obtusata = Platanthera obtusata Habenaria orbiculata = Platanthera orbiculata Habenaria peramoena = Platanthera peramoena Habenaria psycodes = Platanthera psycodes Habenaria purpurascens = Platanthera purpurascens Habenaria saccata = Platanthera stricta Habenaria sparsiflora = Platanthera sparsiflora
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Habenaria stricta = Platanthera stricta Habenaria straminea = Pseudorchis straminea Habenaria tipuloides var. behringiana = Platanthera tipuloides var. behringiana Habenaria unalascensis = Piperia unalascensis Habenaria unalascensis var. elata = Piperia elongata Habenaria unalascensis var. maritima = Piperia elegans Habenaria viridis = Coeloglossum viride var. viride Habenaria viridis var. bracteata = Coeloglossum viride var. virescens Habenaria zothecina = Platanthera zothecina Hybrids: Habenaria xandrewsii = Platanthera xandrewsii Habenaria xbicolor = Platanthera xbicolor Habenaria xcanbyi = Platanthera xcanbyi Habenaria xmedia = Platanthera xmedia Those species formerly included in the genus Spiranthes and most often found within that species in the majority of the literature. Spiranthes adnata = Pelexia adnata Spiranthes cinnabarina = Dichromanthus cinnabarinus Spiranthes costaricensis = Beloglottis costaricensis Spiranthes cranichoides = Cyclopogon cranichoides Spiranthes durangensis = Deiregyne durangensis Spiranthes elata = Cyclopogon elatus Spiranthes lanceolata = Sacoila lanceolata var. lanceolata Spiranthes lanceolata var. paludicola = Sacoila lanceolata var. paludicola Spiranthes michuacana = Stenorrhynchos michuacanum Spiranthes orchioides = Sacoila lanceolata var. lanceolata
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Spiranthes parasitica = Schiedeella fauci-sanguinea Spiranthes polyanthus = Mesadenus polyanthus Other genera often used Stenorrhynchos lanceolatum = Sacoila lanceolata var. lanceolata Beadlea cranichoides = Cyclopogon cranichoides Beadlea elata = Cyclopogon elatus
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LOOKING FORWARD
JUNE 1998
Historical Orchid Collections from Brooklyn, New
York
Amerorchis rotundifolia forma lineata
Trifling with Triphora and Silly Ciliaris
The Growing Season
New Chromosome Number Determinations in Platanthera
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5th
ANNUAL NORTH AMERICAN
NATIVE ORCHID CONFERENCE Lake Itasca State Park, Minnesota
July 8, 9, 10 & 11, 1998
We will begin at noon on July 8th and continue with speakers'
meetings and a wide variety of programs and workshops on July 9th•
Field trips on the 10th & 11
th present an opportunity to see a diversity
of native orchids in full flower. The two specialties of the conference
will be
Malaxis paludosa BOG
ADDER'S-MOUTH and a special trip to the international boundary in Manitoba to see
Platanthera praeclara WESTERN
PRAIRIE FRINGED ORCHIS in one of the largest stands known-
in 1996 over 20,000 flowering stems were seen
Speakers include:
Welby Smith, author, Orchids of Minnesota Bill
Steele, Spangle Creek Labs Larry Zettler,
Illinois College Lorne Heshka, Orchids of
Manitoba Dianne Plunkett, photographing
orchids Mark Larocque, Piperia mysteries
Paul Martin Brown, Color Variation and Form
Margaret From, Platanthera praeclara Nancy Cowden,
Platanthera ciliaris complex