Hodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 87 January 1985 No. 849 Che New England Botanical Club, Inc. Botanical Museum, Oxford Street, Cambridge, Massachusetts 02138 Conducted and published for the Club, by NORTON H. NICKERSON, Editor-in-Chief DAVID S. BARRINGTON RICHARD A. FRALICK A. LINN BOGLE GERALD J. GASTONY WILLIAM D. COUNTRYMAN MICHAEL W. LEFOR GARRETT E, CROW NORTON G. MILLER ROBERT T. WILCE RHODORA.—Published four times a year, in January, April, July, and October. A quarterly journal of botany, devoted primarily to the flora of North America. Price $20.00 per year, net, postpaid, in funds payable at par in the United States currency at Boston. Some back volumes and single copies are available. Information and prices will be furnished upon request. Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to RHODORA, Botanical Museum, Oxford Street, Cambridge, Mass. 02138. 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Address manuscripts and proofs to: Joan Y. Nickerson Managing Editor, RHODORA Phippen-LaCroix Herbarium, Dept. of Biology Tufts University Medford, Mass. 02155 Second Class Postage Paid at Boston, Mass. PRINTED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS Cover illustration Cypripedium reginae Walt., the Showy Lady’s-slipper, was presumed until recently to be extirpated from New Hampshire; five sites for this exquisite orchid are now known. Original artwork by Tess Feltes, Illustrator. Tbhodora (ISSN 0035-4902) JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 87 January 1985 No. 849 THE ORCHIDS OF NEW HAMPSHIRE'!, FRANCES E. BRACKLEY ABSTRACT The distribution, pollination ecology, and habitats of representatives of the Orchi- daceae which occur naturally in New Hampshire, USA are discussed. The informa- tion was compiled from historic records and new field studies. Key Words: Orchids, floristics, pollination, New Hampshire TABLE OF CONTENTS OF GENERA Arethusa CT GEOR eS), PEs a ane 82 Calopogon: .& «04 @ be 3 me Gee Bele 84 Calypso-- 3: «. % @ 3) 2: -4> > SRA ees Coeloglossum: «-%-3 & « a 42-386 Sos) 20 Corallorhiza G6 @ a We Re Ses Be eee 97 Cypripedium . .... 2. 2. ew we 5 Epipactis se Se Ges ce. we &> ak “Ge Pee eee Galearis. os: 4. 2s -« a 3: «&. < 4 canUe Yea 17 OORVEE: 2s 4 wei o & ee Gee we Isotria i gt ge ee ce et So. SCR 74 iparis’ “4:3 % ss “h: te Copyright, 1985 by F. E. Brackley 2 Rhodora [Vol. 87 INTRODUCTION New Hampshire is located in the northeastern United States between Lat. 42° 15’ N and Lat. 45° 1S’ N. It encompasses 9,304 square miles, over 85% of which is presently forested. New Hamp- shire is known as the “Granite State”, an appropriate nickname, as one-third of all the state’s rock is granite or closely related igneous rock. The landscape and soils of the state reflect effects of total glacia- tion experienced during the Pleistocene. Evidence of glacial activity is especially noticeable in the rugged White Mountains which occupy the central portion of the state and include the tallest peak in the northeast, Mount Washington (6,280 ft.). From the 22 miles of seacoast on its southeastern border to the boreal forest adjacent to Canada, New Hampshire supports a diver- sity of habitats. New Hampshire’s flora consists of species with many different geographical affinities, and several orchids reach their northern or southern limits within the state. New Hampshire is one of the fastest-growing states in terms of population; as a result, wildlife habitats are being altered or de- stroyed at an increasing rate. This account of the Orchidaceae records many extant locations which will surely disappear because of human activity. In the preparation of this work, every attempt was made to iden- tify species carefully and to use correct nomenclature according to the rules of the /nternational Code of Botanical Nomenclature (Sta- fleu er al., 1978). As often as possible the original description of each taxon was studied and, in some cases, the type specimen examined. Not all synonyms of each plant are listed, but only those that were, at one time, in general enough use to be commonly encountered. Plant descriptions, unless otherwise indicated, are based upon plants that I have collected and/or examined from New Hampshire. In cases where the number of specimens was few, plants from Maine and Vermont were also examined. The distributions and flowering dates of New Hampshire orchids were gathered from collections, observations, and the literature. Specimens from the following her- baria were studied: AMES, HNH, KESC, MASS, MAINE, NHA, NEBC, VT, and YU. Voucher specimens of my collection are depos- ited in NHA. 1985] Brackley — Orchids of New Hampshire 3 All the known chromosome numbers of taxa occurring in New Hampshire are listed in the discussions of individual species. Unless otherwise indicated, all chromosome counts are taken from A Chromosome Atlas of Flowering Plants, Darlington and Wylie, 1956. THE ORCHIDS OF NEW HAMPSHIRE Key to Genera 1. Lip saccate or slipper-shaped ......... 0.0.00 ccc eeeaeeee (2) 2. Lip slipper-shaped; flowers 1-3, pink, pink and white, or yel- VOW CARE WN onde nw 3 wi es dee. (3) 3. Plant with 2 or more pubescent leaves clasping stem; edges of lip opening in-rolled .............. 1. Cypripedium 3. Plant with one basal petiolate leaf; edges of lip opening turned out and forming a white apron.... 16. Calypso 2. Lip saccate; flowers several to many, whitish to mauve .... cian ARE Eta Maret a wes cea eae aa (4) 4. Leaves alternate in a cluster at the base of the stem; flowers WIRE 605 pcre hae xo ace eee ees 8. Goodyera 4. Leaves alternate along the stem; flowers whitish to mauve eer es re ee ee ee ee re re 5. Epipactis 1. Lip variously shaped but not saccate or slipper-shaped ... (5) 5. Leaf, when present solitary or undeveloped, sheath-like, or absent; stems with or without sheaths .............. (6) G. Leal undeveloped or Aleent o.c<5d cen eee eee (wa) 7. Inflorescence from a corralline-like rhizome; flowers many, variously colored, sepals and petals spreading Cty Shak eke ARN AwESEA Ss ee 17. Corallorhiza 7. Inflorescence from a corm; flowers usually solitary, rose- pink (rarely white), sepals and petals forming a loose hood over the column ............... 12. Arethusa G.. Leal well developed: ic is5665 00 hc be ddd ese ekiass (8) 8. Leaf-blade ovate to obovate; flowers greenish-white or MENOWHEN cos ohGe 44h a 1a ene ee eee eee s (9) 9; Lap SpUrred 45504 044545%% 4. Platanthera (Habenaria) 9. Lip without a spur ................... 14. Malaxis Rhodora [Vol. 87 8. Leaf-blade linear to lanceolate; flowers pink to magenta CPANEL WO vec acta e ee wtih ake eas (10) 10. Lip in lowermost position (resupinate)............ Pie ee sa eee etna ee ed dha Cee eanaes 11. Pogonia 10. Lip in uppermost position (non-resupinate)........ Gertans Reewees 22 Uae eee ee ek ess 13. Calopogon 5. Leaves two or more ........ cc eee ee eee eee eens (11) , OAVOS TWO cy oud hee 6s Sah ee nee hee ees (12) 12. Leaves located near the middle of the stem; flowers small, greenish to purplish ............. 6. Listera $2, MOBVGs- Dasa x caesess nth cote e han o ee osea deers (13) 13. Leaves broadly elliptic to orbicular, lying flat on the BIOUD G4 ae ca peas’: 4. Platanthera (Habenaria) 13. Leaves oblong-elliptic, ascending ............. (14) 14. Stems without foliaceous bracts; flowers green, lat- eral petals reflexed and thread-like .......... ee ee ee ree eT ee eee er eT 15. Liparis 14. Stems with foliaceous bracts; lateral petals conni- vent with sepals to form a hood ............ ore ee eee nee 2. Galearis (Orchis) Leaves MOre Than 1WOss66-2564559%0s ee eeiSeeaed (15) 15S. Leaves at the summit of the stem, appearing whorled Saree ore eee See eT ore er eT ee ee ee 10. Isotria 1S. Leaves scattered along the stem or appearing basal eT Te TET CeCe eT Te eT ere Cae (16) 16. Leaves appearing basal; plant slender with small white spiralled flowers ............0000 9. Spiranthes iO, LOAVES Cau 06 ox div cia soe iene eked heeeee tel (17) 17. Leaves ovate; flowers nodding, pink or white, with- Out spurs; tiny delicate flowers of Beech WOOGS 545445 404 eee baueens 7. Triphora 17. Leaves clasping stem, linear-lanceolate to oblanceo- late, becoming reduced to bracts; flowers not nodding, with spurs; plants of various habitats eee TL ee eee (18) 18. Lip with long slender spur ...............0.. Cup a ee x ad aes Gen 4. Platanthera (Habenaria) 18. Lip with short sac-like spur .............000. ee ee eee 3. Coeloglossum (Habenaria) 1985] Brackley — Orchids of New Hampshire : GENERA AND THEIR INDIVIDUAL TAXA 1. Cypripedium L. Species Plantarum 2:951.1753. The genus Cypripedium was named by Linnaeus who based his description on the Yellow Lady’s-slipper, C. ca/ceolus. The generic name is from two Greek words, Avpris and podion, that have been Latinized [incorrectly, according to Schultes and Pease (1963)]. A traditional translation is “Venus’s shoe” from which the common name “Lady’s-slipper” or “Lady-slipper” is derived. The Lady’s-slippers are the largest and most conspicuous of our native orchids. Flowers may be borne singly, or there may be sev- eral on one stalk. The blossoms are yellow, pink and/ or white. They are composed of three sepals, two of which are usually fused together, and three free petals. Two of the petals are usually narrow and spreading, while the third petal, the labellum, or lip. is expanded into an inflated pouch (the “slipper”). The pistil and stamens are fused together into a compound struc- ture known as the column. The two fertile stamens, which are later- ally located on the column, produce the sticky masses of pollen grains. Pollen in this genus is not organized into pollinia but is loose, granular, and embedded in a viscid secretion (Garay, 1960). On the front of the column, the large sterile stamen, the “stami- node,” is found. Cypripediums possess fibrous roots that have a characteristic acrid odor. The stems grow annually from subterranean rhizomes and bear two or more plicate, conspicuously ribbed leaves. Plants of most species in this genus are covered with glandular hairs which occasionally can cause an itchy rash. In the past, a drug was extracted from the roots of Crpripedium plants for use in treatment of nerve disorders (Hocking, 1955). This medicinal use formed the basis for the old common name, “nerve root.” World-wide, there are 30-50 species of Cypripedium, all found in the Northern Hemisphere. The genus is cytologically uniform in those species for which chromosome numbers have been deter- mined. Diploid counts recorded by Stoutamire (1967) are 2n = 20 or 22. Both natural and artificial hybrids have been reported within Cypripedium (Curtis, 1932). Eleven different species, several varie- ties, and a few hybrids occur in North America. Of these, five species grow wild in New Hampshire. 6 Rhodora [Vol. 87 The Cypripedium flower displays bright color(s), fragrance, and a landing platform, attributes that are most attractive to bees. All of these attributes suggest the presence of nectar, a food source for insects. Since no nectar is actually present, the attractions are deceitful, and the blossom serves as a trap for the deluded insect. Once the visitor enters through the opening in the pouch it is pre- vented from retreating by the involute edges and downward- pointing hairs. To exit from the flower the insect must crawl up through a narrow channel toward the base of the lip. There, two small openings, one under each anther, provide a means of egress. As the pollinator approaches one of the openings, its back must brush against the column. The rough column scrapes off any pollen that the insect may earlier have picked up from another flower. The insect must next squeeze past an anther. In doing so, it dislodges a fresh pollen mass which then becomes firmly cemented to the insect’s back. The insect finally attains freedom via one of the small apertures, carrying with it the pollen necessary to fertilize another Lady’s-slipper, thus effecting cross-pollination. Adaptation to an efficient system of outcrossing means that size of the insect visitor must be compatible with the dimensions of the entire escape route through the flower. L. A. Nilsson (1979) found that some bees are able to lift themselves back out through the entrance of smaller slippers, but an oversized insect visitor may become imprisoned within the slipper and die (Arthur, 1962). When the ovules are fertilized an elongate capsule is formed. Populations of Lady’s-slippers usually produce many flowers but very few fruits. Bingham (1939) and Correll (1950) have suggested as a possible explanation for the low percentage of seed pod forma- tion that there may be a lack of locally available pollinators to transmit the pollen. They also noted that during the lag-period between pollination and fertilization (up to four weeks) the plants are vulnerable to various factors which might prevent fertilization and cause the capsule to abort. Case (1964a) noted that late frosts are a factor in preventing some seed capsules from ripening in the area of the western Great Lakes. A ripened Cypripedium capsule can contain as many as 54,180 seeds (Stoutamire, 1964). However, the time period from seed ger- mination to a mature flowering plant encompasses many years, and 1985] Brackley — Orchids of New Hampshire 7 seedling mortality rate is very high (Curtis, 1943). In general, wild Cypripediums in New Hampshire do not take well to transplanta- tion. Even those plants which at first appear to survive transplanta- tion die after a few seasons and meantime may not reproduce. Clearly, Cypripediums are best left and enjoyed in their natural environments. Further discussion of the North American Yellow Lady’s-slippers is in order. They are generally recognized as varieties of Cypripe- dium calceolus, a Eurasian species described by Linnaeus (1753). His epithet ca/ceol/us means “little shoe.” Although the two taxa of Yellow Lady’s-slippers occurring in New Hampshire originally were described as separate species, most manuals treat them as C. calceo- lus variety pubescens (Large Yellow Lady’s-slipper) and C. calceo- lus variety parviflorum (Small Yellow Lady’s-slipper). Another taxon, C. calceolus variety planipetalum, found in Newfoundland and Quebec, is considered to be most like the Eurasian C. calceolus var. calceolus (Fernald, 1950a). Willdenow described the larger North American Yellow Lady’s-slipper as C. pubescens in 1804. In 1930 this taxon was transferred to C. calceolus as variety pubescens by D. S. Correll. The Small Yellow Lady’s-slipper was first described as Cypripe- dium parviflorum by R. A. Salisbury based upon a specimen from Virginia. In 1946 Fernald treated it at the varietal level as C. calceo- lus variety parviflorum. Many workers have speculated on the relationship between Cypripedium pubescens and C. parviflorum. llitis (1965) listed them as a vicarious species pair that probably diverged during Pleis- tocene glaciation. He theorized that post-glacial migration pro- duced an overlapping of the range of the two races. The resulting hybrids display an intergradation of morphological characters that can be observed today in some populations of Yellow Lady’s- slipper. In New Hampshire the two Yellow Lady’s-slippers appear quite distinct. While the morphological characters of size, colora- tion, pubescence, and overall appearance may be attributed to genotypical response to environmental conditions, these same environmental conditions operate as the primary selective factors in the course of speciation (Sanford, 1974). Sanford also noted that with “small, spatially scattered populations, the development of 8 Rhodora [Vol. 87 slight differences in blooming time becomes an important isolating mechanism.” Mayr (1969) characterized species as groups of interbreeding natural populations that are reproductively isolated. The two races of Yellow Lady’s-slippers in New Hampshire are apparently repro- ductively isolated by flowering time, habitat, and pollinators; they are therefore here recognized as distinct species. Key to Species of Cypripedium in New Hampshire 1. Plants with two basal leaves; flowers solitary; pouch pink or sometimes white, cleft longitudinally down the front ...... ee ere eee Tere ee er Teer ee 1. C. acaule |. Plants with 2 or more leaves borne on the stem above the base; flowers 1-3; pouch yellow, pink or white, often streaked with purple (rarely pure white), not cleft, opening above CCAR va hcew exau 55 Awd eee eth eee Leo oS (2) 2. Lateral sepals separate almost to base; pouch cone-shaped, prolonged downward, white streaked with purple ....... ee rT eee ee ee ee rere Tee errs 2. C. arietinum 2. Lateral sepals fused for entire length, or nearly so; pouch slipper-shaped, erect, yellow or pink (rarely pure white). ... [KEG SE SESES TELLER AL DE EERE TAEO ES SRE TEER EAS B89 (3) 3. Pouch yellow; lateral petals conspicuously twisted, acute at apices, longer than- the Pouch sy x.0s ey cers e neve wees (4) 4. Plants densely glandular-pubescent; sepals and lateral petals greenish-brown or yellow; pouch 3-5 cm long; plants of mesophytic woods, usually flowering in May Peer eT ee ee ne eee rte 3. C. pubescens 4. Plants scarcely pubescent; sepals and lateral petals reddish-brown to purple; pouch 2-2.5 cm, plants of circumneutral or calcareous wetlands, usually flower- [Ne 00 JUNE ces Pacey ers aeeexes 4. C. parviflorum 3. Pouch pink or pink-and-white (rarely pure white); lateral petals straight, white, ovate, shorter than the pouch; plants of circumneutral or calcareous wetlands........ Oe ee ree eee ere ret st re 5. C. reginae 1. Cypripedium acaule Ait., Hort. Kew., ed. 1, 3: 303. 1789. 1985] Brackley — Orchids of New Hampshire 9 The Pink Lady’s-slipper was first described by British botanist and gardener William Aiton from a specimen obtained in North America. This species has also been known as Fissipes acaulis. The literal meaning of the specific epithet acau/e is “stemless.” COMMON NAMES: Pink Lady’s-slipper, Moccasin flower FLOWERING DATES IN NEW HAMPSHIRE: late May through June DescRIPTION: Plant froma short rhizome with thick, fibrous roots having a pungent odor. Leaves two, opposite, basal, elliptic to oblong-elliptic, 1-2.5 cm long, strongly ribbed, and sheathing the flowering stalk. Flower terminating scape, solitary, nodding, sub- tended by an erect bract. Lateral petals and sepals yellowish-green to brownish, often tinged with purple. Lateral sepals joined entirely beneath. Lateral petals narrow, loosely spiraled, inner surfaces pubescent near base. Lip an inflated pouch 4-6 cm long, lightly fragrant, pink (sometimes white in forma a/biflorum) with a longi- tudinal fissure down the front. Edges of opening infolded. Capsule ellipsoid, ribbed, 3-4.5 cm long, held nearly horizontal to ground. CHROMOSOME NUMBER: 2” =20 In late spring and early summer, the woods of New Hampshire are lit up by the large, nodding blossoms of the Pink Lady’s-slipper. It is our most conspicuous and familiar orchid, a delight of the wildflower enthusiast. Cypripedium acaule thrives in the sandy, acidic soil that prevails in most of the state. It commonly grows in open pine or mixed woods, but can be sometimes found in damper, mossy habitats. Occasionally there are years of spectacular blooms. Stuckey (1967) found that unusually large numbers of blossoming plants occurred in areas of Rhode Island 10-15 years after a brush fire. This obser- vation corresponds with the reported average number of years required between seed germination and the production of the first flower in C. acaule (Curtis, 1943). Burning may have a favorable influence in reducing competition for young seedlings and/ or releas- ing necessary nutrients to the soil. In New Hampshire Cypripedium acaule is widespread and can be found in every county. The color of the lip is commonly pink, 10 Rhodora [Vol. 87 although white-lipped plants [forma a/biflorum Rand and Redfield (1894)] become increasingly more frequent in the mountains and the northern areas of the state, even to the point of being the domi- nant form in some populations (Anderson, 1936). Double-flowering forms have been found where the two slippers occur side-by-side or back-to-back on a single, doubly thick stalk. The light, sweet frangrance of the Pink Lady’s-slipper has been found by Stoutamire (1967) to be predominantly produced by the sepals and lateral petals. In the same study it was found that the most common pollinators of this species are bees of the genus Bombus (Apidae). In the White Mountains I have observed many blossoms of C. acau/e with their lips filled with scores of small beetles, identified as Anthobium pothos (Staphylinidae) (W. Morse, personal communication). The beetles appeared to be feed- ing on pollen, but are seemingly too small to serve as effective pollinators. However, Smith (1863) reported that small flower- beetles pollinated the Showy Lady’s-slipper by moving minute par- ticles of pollen from the anther to the stigma. The insects he observed were later identified as Anthobium convexium (Guignard, 1886). A type of self-pollination may be effected by the tiny beetles, but the major cross-pollinator of the Pink Lady’s-slipper is the larger Bombus bee. In any New Hampshire population of Cypripedium acaule, the percentage of plants that set fruit is consistently low, usually less than ten percent. GENERAL DISTRIBUTION: Northeastern North America, Newfound- land west to Alberta, south in the mountains of Georgia and Ten- nessee, west to northern Indiana, northern Illinois and Minnesota. 2. Cypripedium arietinum R. Br., Hort. Kew., ed. 2, 5: 222. 1813. British botanist Robert Brown originally described the Ram’s Head Orchid from a North American specimen. The species has also been known as Criosanthes arietina (R. Br.) House (1905). The epithet arietinum means “of a ram.” COMMON NAMES: Ram's Head Orchid, Ram’s Head Lady’s-slipper FLOWERING DATES IN NEW HAMPSHIRE: May to early June 1985] Brackley — Orchids of New Hampshire 1] DESCRIPTION: Plant small, averaging (in this area) 20 cm tall, from a short rhizome with fibrous, musky-smelling roots. Leaves 3-5, spirally arranged, elliptic-lanceolate, purplish-brownish, lateral sepals and petals narrow, slightly twisted, greenish-purplish. Lip protuberant, prolonged downward, netted with magenta veins. Edges of the circular orifice folded in, covered with a white silky pubescence. Capsule ellipsoid, about 2 cm, prominently ribbed. CHROMOSOME NUMBER: 2n = 20 (Moore, 1973) Looking at the unusually-shaped flower of Cypripedium arieti- num, one can observe an image of the head of a tiny ram in the position for a charge. The Ram’s Head Orchid is the smallest and rarest of all the Lady’s-slippers in New Hampshire. The protuberant lip is “small enough to be put in a child’s thimble” according to Baldwin (1894). Cypripedium arietinum is so inconspicuous as to be difficult to discern fron its surroundings even in blossom. In bud it is almost impossible to find. | have walked by several flowering plants before “discovering” the tiny orchid in a well-known Maine location. Cypripedium arietinum occurs mostly in wet Northern White Cedar (Thuja occidentalis) woods but is also found on well-drained, ledgy slopes under deciduous trees. Cool soils and partial shade are a necessary prerequisite for this species. Although the Ram’s Head Orchid may be locally abundant in some places, none of the New Hampshire stations has ever yielded more than a few plants. There are voucher specimens of C. arietinum from only eleven towns in New Hampshire, and most of these specimens were collected well over 50 years ago. Cypripedium arietinum is listed as “declining” and “vulnerable” in New Hampshire by Storks and Crow (1978), and as “threatened” or “endangered” in New England by Crow er a/. (1981). Despite the scarcity of C. arietinum in New England there are apparently suffi- cient populations of this species on a national scale to warrant its being dropped from consideration as an Endangered or Threatened species of the United States (Federal Register, Vol. 45, No. 242, Monday, December 15, 1980). At present, there is only a single known New Hampshire station for this species. However, I am reasonably confident that more of these dainty orchids are living and blooming in secret haunts within the state. 12 Rhodora [Vol. 87 The Ram’s Head Orchid is considered structurally primitive because there is no fusion of the lateral sepals as in other Cypripe- dium species. Since the flower is so small, only a correspondingly small insect can be an effective pollinator. Stoutamire (1967) observed two bees of the genus Dialictus (Halictidae) visit and carry off pollen masses of C. arietinum on their thoraxes. Van der Pijl and Dodson (1966) list another small bee, Megachile sp. (Megachi- lidae) as a pollinator. Though short lived, the flowers of the Ram’s Head Orchid are not literally ephemeral as once believed. If polli- nation occurs, the upper sepal drops down over the lip, sealing off the entrance way to further insect visitors (Case, 1964a). GENERAL DISTRIBUTION: Quebec west to Manitoba, south to Maine, New Hampshire, Vermont, and New York, west to Michi- gan, Wisconsin, n. Illinois to Minnesota. Cypripedium plectro- chilon of western China may be synonymous with C. arietinum (Leur, 1975). 3. Cypripedium pubescens Willd., Sp. Pl. 4: 143. 1805, a] The name pubescens means “downy,’ fine hairs that cover this plant. in reference to the many COMMON NAMES: Large Yellow Lady’s-slipper, Downy Yellow Lady’s-slipper, Whippoorwill Shoe FLOWERING DATES IN NEW HAMPSHIRE: May to early June DESCRIPTION: | Plant robust, densely glandular-pubescent, 10-70 cm tall, from a cylindrical rhizome with fibrous, musky-smelling roots. Leaves 3-5, elliptic to ovate-lanceolate, strongly ribbed, bright green, sheathing the stem. Flowers | or 2, terminating the stalk, subtended by a large foliaceous bract that usually exceeds the flower in height. Lateral sepals united beneath lip, almost to apex. Dorsal sepal largest, 3-8 cm long, slightly spiralled with undulating margins. All sepals yellowish-green with streaks of brown. Lateral petals narrow, acuminate, loosely twisted to spiralled, yellowish- green. Lip ovate or “slipper-shaped,” usually pale yellow, although all shades of yellow occur. Edges of round opening turned under, inner pouch with reddish-purple spots. Staminode triangular, yel- low spotted with madder. Capsule ellipsoid, ribbed, almost erect. 1985] Brackley — Orchids of New Hampshire 13 CHROMOSOME NUMBER: 2n = 20 In New Hampshire the Large Yellow Lady’s-slipper is most often found in rich deciduous woods along the edges of spring run-off streams. This species prefers a substrate nearly neutral in pH, one reason why it is listed as “infrequent” (Storks and Crow, 1978) here in New Hampshire, the “Granite State.” Curtis (1943) found plants growing in soils which ranged in reactions from pH 5.7 to pH 7.0, but reported that seedlings were most abundant in moist soils that ranged fron pH 6.9 to pH 7.2. Curtis-also noted that the average time period between seed germination and flowering in Cypripe- dium pubescens is 12 years. It has been claimed that many populations of the Large Yellow Lady’s-slipper in New Hampshire have been commercially exploited by persons digging up and selling the plants. For whatever reasons, the records indicate that the numbers of Large Lady’s-slipper pppu- lations have declined in the state; there are only 12 stations docu- mented by specimens. The pollination biology of Cypripedium pubescens has been stud- ied by Robertson (1928) who observed two long-tongued bees of the genus Ceratina (Anthophoridae) visiting the flowers. In Oak- land County, Michigan, Stoutamire (1967) also observed a male Ceratina calcarata enter a slipper and subsequently exit with a pollen mass on its thorax. As with all Cypripediums, percentage of seed set in the Large Yellow Lady’s-slipper is small and propagation is assumed to be largely vegetative. GENERAL DISTRIBUTION: Nova Scotia to Minnesota, south to New England, along the mountains to Georgia, Alabama, Tennessee, and Missouri. 4. Cypripedium parviflorum Salis. Trans. Linn. Soc. I: 77. 1791. The name parviflorum means “small flower.” COMMON NAME: Small Yellow Lady’s-slipper FLOWERING DATES IN NEW HAMPSHIRE: June DESCRIPTION: Plant slight and dainty, 35 cm or less, not as leafy as the preceding, scarcely pubescent. From a small rhizome with fibrous roots that have a pungent odor. Leaves small, 3—5, (usually 4 14 Rhodora [Vol. 87 here), bluish green, strongly ribbed, ovate-lanceolate, sheathing the stem. Floral bracts small, usually not exceeding the flower in height. Flowers usually solitary, strongly fragrant, borne high above the leaves and terminating the stem. Sepals and lateral petals madder- purple. Lateral sepals united almost to the apex. Dorsal sepal 2-4 cm long with undulating edges. Lateral petals lanceolate, tightly spiralled, spreading. Lip small, saccate, bright yellow with purple stripes or spots. Opening circular with inrolled edges, claret spots within. Staminode triangular, bright yellow, spotted with purple. Capsule ellipsoid, almost erect. CHROMOSOME NUMBER: 2n =20 Unlike the almost scentless Large Yellow Lady’s-slipper, the dainty, wax-like blossoms of the Small Yellow Lady’s-slipper pos- sess a strong, sweet fragrance. This species also prefers a wetter, more calcareous habitat and is often found growing in association with Thuja occidentalis. There are only eight stations in New Hampshire where specimens designated “var. parviflorum” have been collected. However, this species may be even more infrequent than the records indicate. Some determinations are mistakes based on confusion over changes in nomenclature, and there are a few specimens which are actually small individuals of the polymorphic Cypripedium pubescens. Van der Pijl and Dodson (1966) listed the bee Ceratina sp. (Anthophoridae) to be the pollinator of Cypripedium parviflorum as observed by Robertson. This information could be misleading if it is interpreted as referring to the Small Yellow Lady’s-slipper. Robertson (1928) listed the long-tongued bee, Ceratina sp., as a visitor to “Cypripedium parviflorum pubescens,” a synonymous name in use at the time for the Large Yellow Lady’s-slipper (Robin- son and Fernald, 1908). Robertson's published blooming dates (May 7-30) correspond more closely to the flowering time of C. pubescens than to the later blooming C. parviflorum. Stoutamire (1967) lists no pollinator specifically for Cypripedium parviflorum. | have observed several small bees visiting the blossom of the Small Yellow Lady’s-slipper. One bee, Lasioglossum zona- tum (Halictidae) was captured with a pollen-smear on its thorax. The smaller size of the pouch in this species indicates that for effec- tive cross pollination to occur, the pollen vector should be smaller 1985] Brackley — Orchids of New Hampshire 15 than the insects observed visiting C. pubescens. Indeed, one visitor to the blossom made repeated attempts to enter the pouch but couldn’t fit through the opening. This larger bee was identified as Megachile melanophoea melanophoea (Megachilidae) (J. Burger, personal communication). GENERAL DISTRIBUTION: Restricted to the northeast. Newfoundland west to Minnesota, south to New Jersey, Pennsylvania and a few isolated stations in the Appalachian Mountains (Case, 1964a). 5. Cypripedium reginae Walt. Fl. Carol. 222. 1788. American botanist Thomas Walter first described the Showy Lady’s-slipper in Flora Caroliniana based on a type specimen from Carolina. This species has also been known as Cypripedium specta- bile Salis. and C. hirsutum Miller. All three names are appropriately descriptive for this beautiful plant. Reginae means “of the queen,” spectabile means “showy,” and hirsutum translates as “hairy.” COMMON NAMES: Showy Lady’s-slipper, Queen Lady’s-slipper, Big Pink and White FLOWERING DATES IN NEW HAMPSHIRE: Mid-June to mid-July DESCRIPTION: Plant froma black cyclindrical rhizome with fibrous roots. Leaves 3-6, ovate-lanceolate, strongly ribbed and hirsute, closely sheathing the stem, bright green. Inflorescence 1-2 (occa- sionally more) large flowers on pedicels arising from axils of erect bracts. Lateral petals and sepals white, ovate, shorter than the lip, spreading. Lateral sepals united entirely. Lip inflated, spherical with infolded edges around the circular opening, deep rose with white markings. Staminode cordate-ovate, white with yellow spots. Capsule ellipsoid, almost erect. CHROMOSOME NUMBER: 2n = 20. Truly the queen of the Lady’s-slippers, Cypripedium reginae is the largest (up to 90 cm) and most spectacular of our native orchids. Since it usually occurs in large colonies, observing the Showy Lady’s-slipper in flower affords an unforgettable experience. The large, lightly fragrant, rose and white blossoms are framed advan- tageously by a background of soft green foliage. Morris and Eames (1929) stated that not even gorgeously colored tropical orchids can 16 Rhodora [Vol. 87 compare with the beauty of this species “for they lack their terrestrial sister’s dower of rich green foliage.” Cypripedium reginae requires a wetter habitat than the other Lady’s-slippers of the region. This species prefers neutral to alkaline soil conditions, but can grow in sphagnum if the roots are able to penetrate to a lower, sweeter soil layer (Case, 1964a). Seedlings, especially those that have developed close to the soil surface, have a very high mortality rate. Curtis (1943) has found that seedlings of the Showy Lady’s-slipper require a three-year growing period before the first green leaf appears above the ground, and that the first flowers may take as long as sixteen years after seed germina- tion to appear. According to Waterman (1949), seeds of this species germinate best when planted deeply (2.5-5 cm). His work in central Michigan suggests that the Showy Lady’s-slipper thrives in those sites where the seeds are pressed into the soil by the feet of deer (Waterman, 1950). Another reported ecological requirement of C. reginae is availability of light. In some localities lumbering may provide the necessary open areas for the plant to prosper. However, Harvais (1980) found in northwestern Ontario that the Showy Lady’s-slipper is more tolerant of shade than it is of low moisture conditions. Historically, only one station for Cypripedium reginae was doc- umented for the state of New Hampshire. The orchid was found growing in Bottomless Pit Bog in Hanover, Grafton County, adja- cent to the Lebanon town line and erroneously assigned to that town on many herbarium labels. The most recent collection of the Showy Lady’s-slipper from that site was made in 1891. In a span of 90 years the site has changed from an open-water bog to a Black Spruce (Picea mariana)-shrub community (J. Poole, personal communication). Because conditions have changed to the extent that the bog no longer supports an orchid population, C. reginae was considered “possibly extinct” for the state (Storks and Crow, 1978). During the course of my field-work, | received information ccn- cerning a possible population of Showy Lady’s-slippers in New Hampshire from Frederic L. Steele, a well-known local botanist. After two days of searching through swamps in the given location | came upon a population of over 200 plants of C. reginae in full flower. The new station is a rather ordinary swamp with Cinnamon 1985] Brackley — Orchids of New Hampshire 17 Fern (Osmunda cinnamomea), Sensitive Fern (Onoclea sensibilis) and Swamp Saxifrage (Saxifraga pensylvanica) growing in abun- dance (Brackley, 1979). Subsequently, I have found two other sites in New Hampshire for the Showy Lady’s-slipper. Both these populations are found in Cedar (Thuja occidentalis) swamps that support other orchids and associated northern elements of the flora as well. Because of the striking beauty of this species there is a danger of exploitation by private and commercial cultivators. In New Hampshire, such a practice could readily cause the demise of this most elegant plant. The report that hirsute stems and foliage of the Showy Lady’s- slipper can cause a dermatitis reaction in some people was sup- ported by a colleague who developed “poison-ivy” on two occasions after accompanying me to a station in northern N. H. where poison- ivy is rarely found. Guignard (1886) observed bees of the genus Megachile (Megachi- lidae) pollinating the Showy Lady’s-slipper. Stoutamire (1967) also observed a medium-sized black bee enter the pouch and quickly force its way out under the anther and carry off a pollen load. In the same study, the fact was noted that although the opening in the labellum allows access to large bees, the exit canal in Cypripedium reginae is smaller than in C. acau/e. This situation suggests that the larger bees of the genus Bombus (Apidae) better “fit” C. acaule. As previously mentioned, Smith (1863) found the small beetle Antho- bium pothos (Staphylinidae) effecting self-pollination in a popula- tion of Showy Lady’s-slippers in Norway, Maine. In Ontario, Arthur (1962) has found that adults of the European Skipper But- terfly Thymelicus lineola (Hesperidae) commonly become trapped within the pouch of the Showy Lady’s-slipper where they eventually starve or are drowned in rainwater. GENERAL DISTRIBUTION: Newfoundland west to Manitoba. south to New England, and southward along the Appalachian Mountains to Tennessee, west through Ohio to Missouri. 2. Galearis Raf., Herb. Raf. 71. 1833. Originally, Linnaeus placed the Showy Orchis in the genus Orchis, a large group of European orchids that provided the basis for the family name, Orchidaceae. Because of morphological differ- 18 Rhodora [Vol. 87 ences (see Vermeulen, 1953) between the European and eastern North American plants, the genus Galearis was segregated from Orchis almost 150 years ago by Rafinesque, the Constantinople- born naturalist who did most of his botanical work in the United States. Because of his eccentricities and his abandonment of the Linnaean principles of taxonomy, Rafinesque’s work was not accepted by botanists of the day. One of his contemporaries de- scribed his prolific writings as “the wild effusians of a literary mad- man” (Humphrey, 1961). In such an atmosphere, Rafinesque’s new genus Galearis was ignored and the Showy Orchis continued to be known by its original name of Orchis spectabilis. When Rydberg revised the genus in 1901, he placed the Showy Orchis in the genus Galeorchis but his generic segregation was not widely accepted. Time has since vindicated some of Rafinesque’s ideas on a natural system of classification and some of his generic segregations, such as Galearis, have been substantiated. Galearis is derived from the Latin word for “helmet,” which fit- tingly describes the hood-like shape made by convergence of the sepals and lateral petals over the column, There are but two species in this genus, the eastern North American G. spectabilis, and an eastern Asian representative, G. cyclochita. 1. Galearis spectabilis (L.) Raf., Herb. Raf. 71. 1833. This species is based upon a type specimen from Virginia. The specific epithet means “showy.” COMMON NAMES: Showy Orchis, Showy Orchid, Preacher-in-the- pulpit FLOWERING DATES IN NEW HAMPSHIRE: May to early June DESCRIPTION: Low plant froma short rhizome with fleshy, narrow roots. Leaves 2, large, basal, sub-opposite, obovate-oblong, nar- rowing into petioles that sheath the 4-5 angled stem. Raceme 2- several flowered, each subtended by a lanceolate floral bract that exceeds the height of the flower. Ovaries thick, sessile, 1-2 cm long. Sepals and lateral petals shell-pink to mauve. Sepals ellipitic, con- cave, connivent to the linear-lanceolate lateral petals, together forming a hood over the column. Lip white (rarely pink in forma Willeyi), entire, dilated, ligulate, having undulating margins and a 1985] Brackley — Orchids of New Hampshire 19 conspicuously broad, club-shaped spur at the base. Capsule ellip- soid, erect, 2 cm long. CHROMOSOME NUMBER: 2n = 42 The Showy Orchis prefers a humus-rich, hardwood forest habitat where it blooms before the leafy canopy fully develops. Although it bears an impressive name, the plant is quite inconspicuous in the dappled light of early spring woods. The shiny basal leaves are often half-concealed under leaf litter, and the nodding, delicately colored flowers are tucked beneath the leafy bracts. Usually, only scattered individual plants are found in flower, accompanied by one or two vegetative plants. Occasionally one finds a small colony of several plants all flowering at once. Blossoming seems to be related to cyclic patterns or specific environmental parameters, as there are “good years” for finding flowering plants; during some growing seasons, vegetative plants predominate and flowering individuals are rarely found. In New Hampshire, the Showy Orchis is generally restricted to the Connecticut River Valley. That area’s alluvial soils tend to be more neutral than the acidic soils which prevail throughout the rest of the state. The Showy Orchis grows with other plants of rich wooded hillsides, such as Maidenhair Fern (Adiantum pedatum), Blue Cohosh (Caulophyllum thalictroides), and Hepaticas (H. acu- tiloba and H. americana). Of the ten historical sites for Galearis spectabilis in New Hamp- shire, only two are outside the Connecticut River Valley. One of these stations, a population discovered by Herbert Sargent in Tuf- tonboro in 1935, seems to have disappeared. A third previously unrecorded population of Showy Orchis in Franklin, Merrimack County was shown to the author in 1982. In New Hampshire the natural habitat of the Showy Orchis occurs within a climax forest community, dominated in this area by Beech (Fagus grandifolia), Sugar Maple (Acer saccharum), and Hemlock (7Tsuga canadensis). In this section of its range, the response of this species to disturbance is highly negative (Auclair, 1972). Unfortunately, the number of undisturbed areas remaining in the eastern deciduous forest is declining rapidly and this loss contributes to the decline of Galearis spectabilis. The species is listed as “vulnerable” by Storks and Crow (1978). The Showy 20 Rhodora [Vol. 87 Orchis apparently has animal predators too, as the last known plant of this species growing in Rhode Island was reported to have been eaten, presumably by a skunk, about 1963 (Stucky, 1967)! Attempts to germinate seeds of the Showy Orchis in the labora- tory have proven unsuccessful. There exists a dormant condition in the ripe seed that is obviously broken in nature, but the means of breaking this dormancy is presently unknown to researchers (Stouta- mire, 1964). Galearis spectabilis is a true food plant for insects (L. A. Nilsson, 1980). It offers rewards to visitors in the form of nectar which is pooled in the broad spur. Robertson (1928) recorded visits by females of Bombus separatus and B. americanorum (Apidae), two species of long-tongued bees. An unusual pink-lipped form of Galearis spectabilis has been named forma Wi/levi for Herman Willey (Seymour, 1970). GENERAL DISTRIBUTION: Eastern North America: New Brunswick south to Georgia and Alabama, west through the southern Great Lake states to Minnesota, and the Mississippi embayment region of Missouri and Arkansas. 3. Coeloglossum Hartman, Handb. Skand. FI. 329. 1820. The genus Coeloglossum was described in 1820 by Hartman based on Coeloglossum viride, originally included in the genus Satvrium by Linnaeus (1753). The name Coeloglossum is derived from the Greek for “hollow tongue,” referring to the saccate spur that characterizes this genus. The genus is currently treated as hav- ing a single species which includes two ecotypes that have been variously interpreted as separate species, subspecies, or varieties; they are here treated on the subspecies level. 1. Coeloglossum viride (L.) Hartman subspecies bracteatum (Mubhl. ex Willd.) Hultén. Coeloglossum viride is a polymorphic species with a widespread distribution throughout the colder regions of the Northern Hemi- sphere. In southern.areas of Europe, Asia, and North America it is confined to cool montane regions (Meusel er a/., 1965; Leur, 1975). In Europe C. viride is commonly known as the “Frog Orchid” or “Die Hohlzunge” (Hollow-tongue) (Summerhayes, 1951; Schrenk, 1985] Brackley — Orchids of New Hampshire 21 1975). The epithet viride is Latin for “green,” the color of the flowers. The North American plant referred to Coeloglossum was origi- nally designated as a distinct species, Orchis bracteata, by Muhlen- berg. He sent a specimen of Long-bracted Orchid from Pennsylvania to Willdenow who published a description of the plant in his Spe- cles Plantarum (1805). In 1813, R. Brown transferred the species to the genus Habenaria. Until recently most American botanists have treated the Long-bracted Orchid as Habenaria viridis var. brac- reata, The conspicuous long floral bracts give this subspecies of C. viride its common name. COMMON NAMES: Long-bracted Orchid, American Frog Orchid. FLOWERING DATES IN NEW HAMPSHIRE: Late may through July. DESCRIPTION: Plant stout, glabrous, from a thickened rhizome, 10-60 cm tall. Roots fleshy. Leaves dark green, 3-5, oblanceolate to lanceolate, clasping the stem. Floral bracts linear, acuminate, much exceeding the length of the flowers. Inflorescence a spike of inconspicuous green flowers. Ovaries sessile, stout. 5-10 mm long. Dorsal sepal ovate, concave. Lateral sepals obliquely oblong. Lateral petals lanceolate. Lip oblong, 3-8 mm long, bi-lobed (sometimes with a small, often obscure tooth in the open sinus), with a thickened median ridge. Spur saccate, greenish-white, approximately 2 mm long. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 40 Terrestrial orchids are known for producing inconspicuous flowers that are often curious-looking rather than aesthetically pleasing. The Long-bracted Orchid is one such species; it is even difficult to determine without a close inspection whether or not the Long-bracted Orchid is blooming. When flowering occurs the sepals and lateral petals barely expand and retain a bud-like form. Only the presence of the green lip extending downward from beneath the hood of sepals and petals verifies that the plant is in full bloom. Ostensibly this species has a wide range of flowering dates, but | have found the Long-bracted Orchid to be an early (May) bloomer, as did Morris and Eames (1929). Microclimatic influences such as o9: Rhodora [Vol. 87 moisture and light availability are apparently important factors in determining flowering time. In New Hampshire, Coe/oglossum viride ssp. bracteatum is most often found in rich deciduous woods. The plants tend to grow singly with only a few scattered individuals at each location. The Long-bracted Orchid is a boreal species and occurs more frequently in the northern areas of the state. Soil reaction studies indicate that this species grows in soils where the pH is between 5.0 and 6.0 (Stuckey, 1967; Wherry, 1918). In addition to the distinctively long floral bracts, the Long- bracted Orchid is characterized by a small, saccate spur, often described in manuals as “scrotoform.” In a study of Ceologlossum viride in the Faroes, Hagerup (1951) found that the wide throat of the nectar-filled spur is covered by a thin membrane. The membrane has only one narrow opening which is hidden behind the thickened base of the lip. In order to partake of the nectar an insect must squeeze its proboscis through the tiny aperture, or penetrate the membrane. Insects with strong biting mandibles, such as beetles, can easily break the thin membrane (Hagerup, 1952). In Finland, Silen (1906) reported that Coe/oglossum viride was pollinated by beetles of the genus Cantharis (Cantharidae). He also observed visits to the flowers by wasps and mosquitoes. | have seen no reports listing pollinators specifically for subspecies bracteatum. The Long-bracted Orchid obviously attracts sufficient pollinators to perpetuate itself; it is one of the most common and widely distributed northern orchids. GENERAL DISTRIBUTION: Boreal North America; south in the mountains to North Carolina and New Mexico; west to Alaska and the Aleutian Islands; Japan and China. 4. Platanthera L. C. M. Rich., Mem. Mus. Paris. 4: 48. 1818. Most species in this genus were originally described as members of the genus Orchis. Although Richard segregated those plants with caudicles of the pollinaria adnate to the column into his genus Platanthera in 1818, most botanists, until very recently, treated these plants as members of the genus Habenaria. Habenaria was described by Willdenow (1805) to accommodate two tropical American orchids with projecting anther caudicles (Stoutamire, 1974). The tropical genus Habenaria as now understood is 1985] Brackley — Orchids of New Hampshire 23 distinguished from Platanthera by its two stigmatic lobes borne on stalks beside or below the nectary opening. Platanthera has a single median stigma above the nectary opening, and is largely composed of plants with temperate distributions (Stoutamire, 1974). Platanthera consists of about 200 species (Leur, 1975). Typically, the base of the lip is spurred, and the length of the spur usually corresponds to the tongue-length of a specific pollinator. In this genus the pollinia are attached to caudicles which are distally united to a sticky disc of cells called the viscidium (L. A. Nilsson, 1978). The entire complex structure has the shape of a drumstick or club, and has been termed a “pollinarium” or a “hemipollinarium” (Dressler, 1981). The two pollinaria are embedded in locules on either side of the opening of the nectary, with the sticky discs of the viscidia facing each other. The distance between the viscidia usually corresponds with the size of the head of a specific pollinating insect. As the insect probes the nectar-containing spur with its proboscis, the viscidia adhere to the pollinator, and the pollinaria are pulled from their locules. At this point, one of the most amazing adaptations for cross-pollination occurs. As the insect flies to another flower to partake of more nectar, the caudicles rotate downward and inward resulting in the pollinaria being in a position to come directly in contact with the stigma of the next visited flower. This phenomenon was studied by Darwin (1877) and by Gray (1862-1863). Most Platanthera species are pollinated by moths and butterflies (Lepidoptera). The name Platanthera is derived from two Greek words signifying “wide-anther.” Key to the Species of Platanthera in New Hampshire 1. Lip fringed or lacerate along margin..................4. (2) 2. Flowers white; lip entire; plants of sphagnum bogs........ ee eee re re ere Serre ee 1. P. blephariglottis 2. Flowers rose, pink, purple or green (rarely white), lip divided into three segments; plants usually not occurring in SONASNUO DOSS «ixckded hee vader Ha Gad eeawd Rx eatia (3) 3. Flowers green; plants of hayfields and roadsides (occasion- BUY WARIS) cen eave cae coon 4 eae ee Cee aS 4. P. lacera 3. Flowers pink, rose or purple (rarely white); plants of wet ditches, marshes and ' WoO0dS: oi466 666 65.0-005454564 (4) 24 Rhodora [Vol. 87 4. Lip typically shallowly fringed; opening of nectary oblong, constricted and appearing as 2 lateral openings ee TOR eT a ee re eer re 2. P. psycodes 4. Lip typically deeply fringed; opening of nectary circular and unobstructed ................3. P. grandiflora be Beh BA cas ag va eee caer eaye sas BELA ews (5) 5. Leaves basal, with or without bracts on stem.......... (6) 6. Leaf solitary, ovate; plants of northern woods .......... Lip uesuen rade bane sare cat pnee eas eee ae a 7. P. obtusata 6. Leaves two, oblong to orbicular, at or near the surface of the ground; plants of mixed or deciduous woods... .(7) 7. Flowers white; lip curved downward; stem with several SANE HAC ova bce nenae vee ee ee us 5.P. orbiculata 7. Flowers greenish-yellow; lip curved upward; stem with One DIScl OF RAKE? o.4:050 fe c5e0ns ok bs 6. P. hookeri 5. Leaves cauline; bracts present............c0ce eee ceees (8) 8. Plant with a single prominent leaf; flowers askew, greenish; spur club-shaped; plant of wet woods and moist ditches i554 R LL EREL REP OSCR GERLE Gao eee ae 8. P. clavellata 8. Plant with two or more prominent leaves; flowers not askew, white, yellowish or greenish; spurs variously shaped but not club-shaped; habitats various ...... (9) 9. Flowers whitish-yellowish; lip bearing a prominent tu- BONE, ee DI aad henna ce eee ee Kas 9. P. flava 9. Flowers white or greenish; lip lacking a tubercle, apex ra 6 na ee an eR, Oe (10) 10. Flowers white, strongly fragrant; lip abruptly dilated Ai. WASC satin eres yea ee IN sees 10. P. dilatata 10. Flowers green, only slightly fragrant; lip gradually widened At DASE v.i444 eure ses 11. P. Ayperborea 1. Platanthera blephariglottis (Willd.) Lindl., Gen. Sp. Orchid. 291. 1835. Willdenow (1805) described this species as Orchis blephariglottis from plants collected in Pennsylvania. This orchid has also been known as Habenaria blephariglottis and Blephariglottis blephari- glottis. The epithet blephariglottis translates as “eyelid-tongue,” in reference to the tongue-shaped fringed lip. 1985] Brackley — Orchids of New Hampshire 25 COMMON NAME: White Fringed Orchid FLOWERING DATES IN NEW HAMPSHIRE: July through August DESCRIPTION: Plant from many fleshy roots, glabrous. Stem leafy, strongly ribbed. Leaves 2-3, sheathing the lower stem, elliptic- lanceolate, somewhat keeled, becoming bracteate above. Inflores- cence a raceme. Floral bracts lanceolate. Flowers snow-white with a cream colored column. Dorsal sepal ovate, concave. Lateral sepals ovate, oblique. Lateral petals linear to oblong, erose at apex. Lip tongue-shaped, fringed at margins. Spur slender, slightly curved. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). Platanthera blephariglottis is an orchid of Black Spruce- Tamarack bogs. It is seldom found on the open mat near the water’s edge where Rose Pogonias, Grass-pinks, and cranberries thrive; its preference is the partial shade of small shrubs and tall grasses. The snow-white spires of the White Fringed Orchid are in bright contrast with the bog greenery. Inspecting an individual flower, one can see that the oval lip is copiously cut along the margins into a feathery fringe. Occasionally plants of this species appear with an unfringed lip, a form that was segregated by Lindley, and graphically named P. holopetala. In Platanthera blephariglottis, the projections of the column in which the pollinaria are embedded are a creamy light-gold color. The tips of the long curved spurs are also of a golden color, indicating the presence of nectar within. Smith and Snow (1976) reported that in Michigan, Platanthera blephariglottis is predominantly pollinated by several species of moths. Insects that were captured in their study carrying pollinaria of the White Fringed Orchid include Darapsa versicolor, Haemor- rhagia thysbe (Sphingidae), Papilio troilus (Papilionideae), and Bombus sp. (Apidae). In the same study, the Spicebush Swallow- tail, Papilio troilus was also found to be a pollinator of the Yellow Fringed Orchid P. ciliaris. This orchid is closely related to the White Fringed Orchid, the major difference being the color of the flowers. When sympatric, the two species occasionally hybridize, most prob- ably through the services of the Spicebush Swallowtail. Although 26 Rhodora [Vol. 87 the Yellow Fringed Orchid was reported from the state of New Hampshire in 1872, it has not been found here since. It is possible that the determination documenting the report was incorrect, as it is difficult to distinguish the two taxa on the basis of dried herbarium material. At the southern end of its range, the White Fringed Orchid is sometimes segregated as var. conspicua on the basis of larger flow- ers and longer spurs (Correll, 1950). Although the White Fringed Orchid is often locally abundant in the state, the species seems to be experiencing a decline in its numbers in New Hampshire. Over an eight-year period, | have observed a drop in the number of individual plants at established locations. Reports of new stations for this orchid are rare. Perhaps the ostensible decline of this species is more apparent than real; there may be many populations of White Fringed Orchids hidden in undiscovered bogs throughout the state. GENERAL DISTRIBUTION: Newfoundland south through New England and the mid-Atlantic states, west to Ontario and Michigan, south to Florida, west to Texas. 2. Platanthera psvcodes (L.) Lindl., Gen. Sp. Orchid. 294. 1835, as to synonymy and name, excluding description. Linnaeus (1753) described this species, which was collected by Peter Kalm in Canada, as Orchis psvcodes. Lindley transferred the taxon to the genus Platanthera, but his accompanying description was of the species that is known today as P. lacera. This orchid is perhaps most well known by American botanists as Habenaria psy- codes. The species name psvcodes is a misspelling of “psychodes,” the Greek word for “butterfly,” perhaps alluding to the shape of the flowers. COMMON NAMES: Small Purple Fringed Orchid, Butterfly Orchid FLOWERING DATES IN NEW HAMPSHIRE: late July to mid-August DeEscRIPTION: Plant glabrous, leafy, from fleshy roots. Leaves 2-5, elliptic to lanceolate, becoming bracteate above. Inflorescence a densely flowered raceme. Floral bracts lanceolate. Flowers mauve or purplish (rarely white), fragrant. Sepals elliptic. Lateral petals ascending, spatulate with denticulate margins. Lip cleft into 3 divi- sions, fringed, reflexed. Spur slender, slightly clavellate. Opening of 1985] Brackley — Orchids of New Hampshire 27 nectary constricted by a projection with the resulting appearance of 2 openings. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Stoutamire, 1974). Platanthera psycodes occurs in New Hampshire in wet grassy areas and along stream sides. This showy species lends its various shades of pinkish-purple flowers to the mid-summer July landscape. In large colonies of the Small Purple Fringed Orchid one can some- times find a rare albino form (forma a/biflora). In our area this species flowers later than the Large Purple Fringed Orchid, P. gran- diflora, with which it is often confused. This confusion arises because the two taxa have been treated by many botanists as varie- ties of a single species. Distinctions between the two taxa in the past have been based largly on size, which, despite the common names, is not a reliable criterion. Stoutamire (1974) did a comparative study of the two orchids and found that in most areas where both occur, the mean height of P. psycodes exceeds that of P. grandiflora. | have never encountered such a situation in New Hampshire perhaps owing to a relatively small number of populations of the Small Purple Fringed Orchid in the state (map 9). An easily observable difference between typical individuals of each species is found in the appearance of the raceme. In P. psycodes the raceme is tightly packed with many small indi- vidual flowers; it has the overall shape of a cylinder. The inflores- cence in P. grandiflora is loosely flowered with fewer, larger blossoms, and is noticeably wider. The tripartite lip of the Small Purple Fringed Orchid is typically reflexed downward while the lip of the Large Purple Fringed Orchid is more widely spreading and curved upward. While these attributes may vary, Stoutamire (1974) found basic morphological differences between the two species that are func- tionally important. He found that the structure of the column and the shape of the nectary are characters that consistentlv separate the two taxa and determine their pollination biology. The nectary open- ing in Platanthera psycodes is oblong in shape but partially con- stricted so as to produce “two lateral openings, one beneath each viscidium” (Stoutamire, 1974), whereas the circular opening of the nectary in P. grandiflora is unobstructed. These seemingly slight differences in floral morphology dictate different pollinators for each species, and thus the two taxa remain reproductively isolated. 28 Rhodora [Vol. 87 S. 1. Smith of Norway, Maine, observed the Hawkmoths Sesia (Haemorrhagia) thysbe and S. (H.) diffinis (Sphingidae), and a Swallowtail Papilio polyxenes asterias (Papilionidae) visit the flow- er of Platanthera psycodes. The insects had “their probosces so encumbered with the pollinia, that it was impossible for them to be coiled up between their palpi” (Smith, 1863). In Michigan, Stouta- mire (1974) also observed the Hawkmoth Haemorrhagia thysbe vis- iting the Small Purple Fringed Orchid, as well as the Skipper Polites mystic (Hesperiidae), each carrying pollinaria of the orchid. Although the scent of the Small Purple Fringed Orchid has been described as sweet (Gray, 1862-1863), | am inclined to agree with Baldwin (1894) who described the odor as “rank.” To Ramsey (1966) the scent of this species is like “fresh calico.” Platanthera psycodes forms natural hybrids with the more wide- spread Ragged Orchid. A hybrid plant was first discovered by M. W. White and A. L. Andrews in a “very wet meadow in Pownal, Vermont, July 22, 1898” (Andrews, 1901}. White (1904) named the hybrid Habenaria Xandrewsii in honor of his friend, Andrews, a well known bryologist, and published it in Niles (1904). The hybrid is intermediate between the two parents and most often occurs in the northern areas of this state. In Nova Scotia, I have seen roadside ditches filled with flowering plants of P. psycodes, P. lacera, and hybrids of every conceivable intermediate form. GENERAL DISTRIBUTION: Newfoundland west through the Great Lakes area to Ontario and Minnesota, south through New England and the mid-Atlantic states to Virginia; continuing southward in the mountains to Georgia. 3. Platanthera grandiflora (Bigelow) Lindl. Gen. Sp. Orchid. 294. 1835. Investigating the nomenclatural history of this taxon is a confus- ing and frustrating ordeal. Fortunately, Stoutamire (1974) has assiduously researched the subject recently and has determined that the correct name for this species is P. grandiflora. He reported that, due to inadequate descriptions and lack of type materials, older names previously ascribed to this taxon must be rejected, thus Bigelow’s (1824) combination of Orchis grandiflora must be taken as the basionym. Bigelow based his description on plants found in 1985] Brackley — Orchids of New Hampshire 29 Massachusetts and Enfield, New Hampshire. The lectotype is de- posited in AMES. American botanists have long known this species as Habenaria fimbriata. The epithet grandiflora means “large-flowered.” COMMON NAME: Large Purple Fringed Orchid FLOWERING DATES IN NEW HAMPSHIRE: late June to early August DESCRIPTION: Plant from fleshy roots, glabrous, leafy. Leaves 2-6, elliptic at base, becoming lanceolate and then bracteate above. Inflorescence a loosely-flowered raceme, approximately 2-4 times longer than broad. Floral bracts lanceolate. Flowers lilac to rose- purple. Sepals ovate-elliptic. Dorsal sepal erect, lateral sepals spreading. Lateral petals oblong with finely dentate margins. Lip cleft into 3 fan-shaped segments, deeply fringed. Spur slender, slightly clavellate. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Stoutamire, 1974). Robert Frost (1964) referred to the Purple Fringed Orchid as “the far sought flower.” It is the largest of our orchids, and one of the most beautiful. Thoreau (1884) stated that Bigelow thought it to be the most beautiful of all the orchids, and added “Is it not significant that some rare and delicate and beautiful flowers should be found only in unfrequented wild swamps?” In New Hampshire, swampy areas are not the only habitats for Platanthera grandiflora. The tall purple wands can often be found along wet, unmowed roadsides, while less robust forms inhabit wet woodlands, sometimes flourish- ing in the darkness of a coniferous forest. It is in open areas that this species attains its greatest size. Ramsey (1966) reported an individ- ual plant of the Large Purple Fringed Orchid a full five feet tall, and Morris and Eames (1929) observed flowers with lips more than | 1/8 inches in width. The loosely flowered racemes of lilac to rose-purple flowers are typically more deeply fringed then in Platanthera psycodes and sug- gested to Baldwin (1894) “a flock of birds struggling to get foot-hold on the same branch.” Such abundance of form and beauty alone would make this species exceptional, but the scent of the flowers elevates the Large Purple Fringed Orchid to a thing of perfection. The fragrance is reminiscent of the sweetest-smelling roses, only it is much nicer. 30 Rhodora [Vol. 87 Flowers of Platanthera grandiflora are adapted for pollination by long-tongued Lepidoptera which transport the pollinaria attached to their compound eyes (Stoutamire, 1974). Van der Pijl and Dod- son (1966), and Moldenke (1949) reported the Large Purple Fringed Orchid as being pollinated by moths. Various species of Swallowtail butterflies have also been reported as visiting the orchid (Stoutam- ire, 1974; Ramsey, 1966). I have also observed these butterflies on the flowers of P. grandiflora. Occasionally, plants of the Large Purple Fringed Orchid occur with narrow and fringeless middle lip divisions. Fernald (1946b) has called such individuals forma menotonsa (with shaved chin). I saw this form, as well as an albino form, in the same swampy area of Rumney, Grafton County, in 1978. GENERAL DISTRIBUTION: Newfoundland west through the Great Lakes area to Ontario and Minnesota, south to Virginia, and con- tinuing southward in the mountains to Tennessee; west to lowa. 4. Platanthera lacera (Michx.) G. Don in Sweet, Hort. Brit. ed. III. 650. 1839. This species was described by Michaux (1803) from plants col- lected in Carolina. It has also been known as Habenaria lacera and Blephariglottis lacera. The epithet /acera is Latin for “torn,” descrip- tive of the deeply fringed lip. COMMON NAMES: Ragged Orchid, Green Fringed Orchid FLOWERING DATES IN NEW HAMPSHIRE: July through mid-August DESCRIPTION: Plant from fleshy roots, glabrous, leafy, up to 60 cm tall. Leaves rigid, 2-5, sheathing lower stem below, oblong to lan- ceolate, becoming bracteate above. Inflorescence a loosely flowered raceme of greenish-white to yellowish flowers. Floral bracts lanceo- late. Dorsal sepal ovate. Lateral sepals oblong, oblique, reflexed. Lateral petals linear-oblong, ascending. Lip deeply tripartite. Lat- eral lobes deeply cut into thread-like fringes. Mid-lobe linear- spatulate, cut about half-way into fringes. Spur slender, slightly clavellate. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Stoutamire, 1974). The Ragged Orchid is a widespread and common midsummer orchid in New Hampshire. This species often grows in open, grassy 1985] Brackley — Orchids of New Hampshire 31 areas where it is hidden in tall grasses. The greenish flowers seem inconspicuous at first, but a closer look reveals a most delicate and lacy blossom. The British horticulturist Robert Sweet described the Ragged Orchid as being “elegantly jagged” (Gibson, 1905). Platanthera lacera is tolerant of a wide variety of habitats. It occurs in both damp and drier habitats, in the open and, sometimes, in the woods. The favored habitat of this species is hayfields where it usually blossoms just before the hay is cut. Several times I have plucked a specimen only minutes before the mower flattened the waving herbage. Morris and Eames (1929), in their discussion of P. lacera, referred to the “deadly swarth of the mower’s scythe,” but I have found that, just as the grasses reappear the following year, so do these hardy orchids. Stoutamire (1974) found the Hawkmoth (Haemorrhagia thysbe (Sphingidae) carrying the pollinaria of both Platanthera lacera and P. psycodes. These insects are probably the agents responsible for producing the hybrid between the two taxa, P. Xandrewsii. Gray (1862-1863) observed that nectar was more plentiful in the spurs of older flowers (“almost full”), especially those individuals which had their pollinaria removed and stigma fertilized. In Newfoundland Platanthera lacera var. terrae-novae, a robust plant with less dissection of the lip, replaces the typical var. /acera (Fernald, 1926b). GENERAL DISTRIBUTION: Newfoundland west to Ontario and Wis- consin, south to Georgia, west to eastern Texas. 5. Platanthera orbiculata (Pursh) Lindl. Gen. Sp. Orchid. 286. 1835. Pursh (1814) originally described this orchid from plants found in shady Beech woods in the mountains of Pennsylvania and Virginia. He noted that the plant was known “in the mountains by the name of Heal-all.” The name orbiculata is Latin for “round,” referring to the two large, round leaves. COMMON NAME: Large Round-leaved Orchid FLOWERING DATES IN NEW HAMPSHIRE: late June through early August DESCRIPTION: Plant from long, fleshy roots, glabrous, scapose. Leaves 2, lying flat on the ground, subopposite, basal, elliptic to 32 Rhodora [Vol. 87 orbicular, shiny blue-green above, silvery beneath. Stem stiffly erect with one or two small bracteate leaves. Inflorescence a loosely flowered raceme. Floral bracts linear-lanceolate. Flowers white, few to 25 or more. Dorsal sepal erect, sub-orbicular. Lateral sepals oblong, obtuse, reflexed. Lateral petals ovate-lanceolate, oblique, spreading. Lip linear-oblong, pendant, recurved. Spur long and slender. Column large with 2 wing-like projections on each side. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). The Large Round-leaved Orchid is a species that, in our region, is confined to woodlands. It is most common in rich deciduous woods but can also be found in mixed or coniferous forests. A favored spot for Platanthera orbiculata is by the base of a large tree. | once found it blossoming in an open, slash-filled area in a cut-over Cedar woods in Coos County. The Large Round-leaved Orchid is not uncommon in the higher, cooler regions of the state, but is rarer in the sou- theastern lowlands. Platanthera orbiculata is a large and impressive plant, even in the vegetative state. The twin leaves, which range in shape from oblong to perfectly round, are sometimes as large as 19 cm in diameter. The shiny, blue-green leaves hug the forest floor closely, but if pried up they reveal a silver surface on their undersides. In a population of Large Round-leaved Orchids, there are usually more vegetative plants than flowering individuals. Exceptionally large plants of the Large Round-leaved Orchid have been described as a separate species, Platanthera macrophylla, which Ames (1906) distinguished on the basis of spur length. Leur (1975) treated the larger plants as P. orbiculata var. macrophylla, while Morris and Eames (1929) referred to them as forma macro- phylla. The majority of plants that I have seen from New Hampshire approach the larger form. Additionally, a dwarfed plant of this species from Newfoundland has been described as var. /ehorsii (Fernald, 1950b). It is apparent that P. orbiculata is exceedingly plastic in regard to size. The large curious-looking flowers have elongated lips and spurs. By night, the white blossoms are conspicuous and lightly fragrant. Night-flying Hawkmoths are lured to the flowers to sip from the nectar-filled spurs. The moth Sphinx drupiferanun (Sphingideae) is 1985] Brackley — Orchids of New Hampshire 33 listed by van der Pijl and Dodson (1966) as a pollinator of Platan- thera orbiculata. Stoutamire (1971) also reported the moths Auto- grapha ampla and Plusia balluca (Noctuidae) to be pollinators of this species. According to Gray (1862-1863) self-fertilization in this species is “out of the question.” The Large Round-leaved Orchid does reproduce vegetatively and it is common to find colonies where the large leaves of several plants overlap. GENERAL DISTRIBUTION: Woodlands from Newfoundland, follow- ing the tree-line to Alaska and the Aleutians, south to Virginia and continuing southward in the mountains to Georgia, west to Washington and Oregon. 6. Platanthera hookeri (Torr.) Lindl., Gen. Sp. Orchid. 286. 1835. This species was described from plants collected in New York, and was named for William Jackson Hooker (1785-1865), an Eng- lish botanist who published, among other works, a flora of northern “British” America (Stafleu, 1967). This species has also been known as P. hookeriana and Habenaria hookeri. COMMON NAME: Hooker's Orchid FLOWERING DATES IN NEW HAMPSHIRE: late May to early July DESCRIPTION: Plant from long, fleshy roots, glabrous, scapose. Leaves 2, basal, sub-opposite, oval to orbicular, light green. Stem olive-colored, naked or with a solitary bract. Inflorescence a loose, elongate raceme of few to many flowers. Floral bracts lanceolate. Flowers yellowish-green, erect. Dorsal sepal ovate to triangular. Lateral sepals oblong-lanceolate, strongly reflexed. Lateral petals linear, falcate, connivent with dorsal sepal. Lip triangular, acute, strongly upcurved. Spur slender. Column with a tubercle above stigma. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). Hooker’s Orchid is one of our early-blooming woodland species with a habitat preference for rich deciduous woods of Sugar Maples and Beech. It is often associated there with other orchids; I have found this orchid growing near the Showy Orchid, reaching its peak bloom a few days after Galearis spectabilis. In New Hampshire this species is widespread but does not occur as often as the Large Round-leaved Orchid. 34 Rhodora [Vol. 87 Superficially, Hooker’s Orchid resembles Platanthera orbiculata but can be distinguished by its smaller leaves that are not typically as flattened as those of the latter. The stem is an olive-green color and is usually naked. The flowers of P. hookeri are quite distinct, being closer to the rachis, yellowish in color, and with a wider, upturned lip. The whole appearance of the flower has been likened to a “gargoyle” (Correll, 1950), or termed “grotesque” (Schrenk, 1977). The most practical description for determining this species 1s given by Morris and Eames (1929) who coined the phrase “Hooker’s hooks” as an “identity-tag.” The pair also stated that the origin of the hooked lip is from the crease of its folding in the bud. In Platanthera hookeri the viscidia are more widely divergent than in our other Platantheras, and an insect would be quite able to steal nectar without coming in contact with the pollinaria if it were not for the upturned lip that prevents a direct approach to the nectary (Gray, 1862-1863). An insect must approach the nectary from the side and in doing so “its face will almost certainly be brought into contact with one of the discs” (Darwin, 1877). Darwin (1877) also reported that Gray observed a butterfly, NMisoniades (Hesperiidae) in Canada with the pollinaria of P. hookeri attached to its eyes. Since the genus Nisoniades is tropical in distribution, only rarely reaching southern Texas, Gray must have seen another Skipper in Canada. GENERAL DISTRIBUTION: Nova Scotia west through the Great Lakes area to lowa, Michigan and Manitoba, south to Pennsylvania. 7. Platanthera obtusata (Banks ex Pursh) Lindl., ssp. obtusata Gen. Orchid. 284. 1835. This species was collected by the British naturalist Sir Joseph Banks, and was described in Pursh’s Flora Americae Septentrionalis in 1814. The type specimen was collected from Hudson Bay in Canada. The name obtusata means “blunt,” and refers to the rounded apex of the solitary leaf. COMMON NAMES: Blunt-leaf Orchid, Small Northern Bog Orchid FLOWERING DATES IN NEW HAMPSHIRE: July to early August DESCRIPTION: Plant from a few fleshy roots, small (usually not over 20 cm tall), glabrous. Leaf solitary (rarely 2), obovate to oblan- ceolate, blunt at apex. Stem naked or with 1-2 bracteate leaves. 1985] Brackley — Orchids of New Hampshire 35 Inflorescence a few-flowered raceme of greenish-white flowers. Floral bracts lanceolate. Dorsal sepal concave, ovate with rounded apex. Lateral sepals lanceolate-elliptic, oblique, reflexed. Lateral petals triangular-lanceolate, apiculate, connivent with dorsal sepal. Lip fleshy, linear-lanceolate, pendant, with a callus in the center of the base that serves to divide the nectary opening. Spur triangulate, elongated, curved. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 126 (ssp. oligantha). The Blunt-leaf Orchid is a small and inconspicuous Platanthera. It is a true plant of the north, being distributed beyond the tree-line into the alpine zone of mountains and tundra of the arctic. This species also occurs in sections of northernmost Eurasia where those plants have been segregated as ssp. oligantha. Subspecies oligantha is a smaller plant with fewer flowers than our ssp. obtusata. Inter- mediates between the two races have been reported from Alaska (Moore, 1980). In eastern North America, the southern limit of the Blunt-leaf Orchid is reached in northern New England. This species is not widespread in New Hampshire as it requires a habitat where midsummer soil temperatures rarely exceed 15° C. (Stoutamire, 1968). Accordingly, P. obtusata is found in the montane locations in the upper-middle region of New Hampshire, and in the cold, swampy woods of the northern part of the state. To find this orchid, One must explore wet, northern woods floored with mosses and humming with insects. Once a plant of P. obtusata is discovered, a quick search of the area will often reveal many more individuals as this species, once established, forms large populations. The whitish- green flowers of the Blunt-leaf Orchid are described as “galeate,” or helmet-shaped, in reference to the hood formed by the convergence of the lateral petals and the dorsal sepal. Although Raup (1930) described this species as possessing scent, I have not been able to detect an odor. It is possible that the flower may become more fragrant toward the end of the day. Platanthera obtusata has the distinction of being one of the few plants known that is pollinated by mosquitoes. I have often observed mosquitoes resting on orchid flowers, but in this case mosquitoes have actually been observed carrying the pollinaria of this species (Raup, 1930; Stoutamire, 1968; Thein, 1969; Thein and Utech, 1970). Although mosquitoes are not well known as pollina- tors, there is evidence that these insects obtain nectar for food in the 36 Rhodora [Vol. 87 higher latitudes (Hocking, 1968). Kevan (1972) reported mosquitoes of the genus Aedes (Culicidae) carrying pollen of the Mountain Avens (Drvas integrifolia) in the high Canadian arctic. Stoutamire (1968) and Thein (1969) found Aedes spp. with the pollinaria of P. obtusata in Michigan and Wisconsin respectively. The callus on the base of the lip of this orchid which divides the nectary opening makes it necessary for the insects to approach the nectary from the side. Mosquitoes probing the nectary come in contact with a viscidium which attaches the pollinarium to the insect’s eye. Moths of the genus Xanthorhoe (Geometridae) have also been observed on the flowers of the Blunt-leaf Orchid, and with the pollinaria attached to their eyes (Stoutamire, 1968; Thein and Utech, 1970). GENERAL DISTRIBUTION: Woodlands, Labrador and Newfound- land, west to the Aleutians, south to northern New England, west through the Great Lakes area, south in the western mountains to Colorado; Norway, Sweden, northern parts of central and eastern Asia. 8. Platanthera clavellata (Michx.) Leur, Native Orchids Fla. 148. 1972. Michaux (1803) named this species Orchis clavellata, which he described from plants collected in Carolina. At one time it was called Habenaria tridentata for the three small teeth at the apex of the lip. The epithet c/avellata means “little club,” referring to the club-shaped spur. COMMON NAMES: Little Club-spur Orchid, Small Green Wood Orchid FLOWERING DATES IN NEW HAMPSHIRE: July through August DESCRIPTION: Plant glabrous, small (S-35 cm tall), from slender, fleshy roots. Stem angled, with several linear-lanceolate bracts and one lower, well-developed leaf, oblong to oblanceolate. Inflores- cence a few- to many-flowered raceme. Floral bracts lanceolate. Flowers whitish to yellowish-green, askew. Sepals ovate, obtuse. Petals ovate, oblique, obtuse. Lip oblong-cuneate, truncate, with 3 obscure notches at apex. Spur slender, clavellate, curved. Capsule ellipsoid, ascending. 1985] Brackley — Orchids of New Hampshire 37 CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). One of our most common orchids, P. clavellata is distinct; no other orchid in our flora carries its flowers at such an angle (“askew”), a feature that is seldom captured in drawings of this species. In New Hampshire, the Little Club-spur Orchid is wide- spread and common. One reason for its abundance is that it can occur in a wide variety of habitats. | have found this orchid in acidic roadside seeps that later yielded prodigious numbers of Nodding Ladies’-tresses (Spiranthes cernua), and in boggy open habitats in the company of Rose Pogonias (Pogonia ophioglossoides). \n deeply shaded woods the Blunt-leaf Orchid appeared with the fruit- ing Early Coral-root (Corallorhiza trifida), and the flowering Large Purple Fringed Orchid (Platanthera grandiflora). In Platanthera clavellata, there is usually only one well-developed leaf-blade near the base of the plant, and 2-5 small bracteate leaves higher on the stem. The flowers seem to be of a more whitish- yellowish color when growing in the open, and more of a greenish hue under shade. The broad wedge-shaped lip has three small notches at the apex. The long curved spur widens at the end into the club-shape that gave this orchid its name. Gray (1882-1883) observed that packets of pollen in Platanthera clavellata often detach themselves from the pollinaria and fertilize the stigma before the flower opens. Because the Little Club-spur Orchid is self-fertilizing, almost all of the flowers form fruit. No doubt, this reproductive strategy is partially responsible for the abundance of P. clavellata in New Hampshire. GENERAL DISTRIBUTION: Newfoundland west through the Great Lakes area to Ontario and Minnesota, south to northern Florida, west to Texas. 9. Platanthera flava (L.) Lindl. Gen. Sp. Orchid. 293. 1835. From plants collected in Virginia, Linnaeus described this species as Orchis flava. \t has had an interesting history of nomenclature, often being confused with Coe/oglossum viride. Some of the many names by which it has been known include Habenaria flava and P. herbiola. Some authors (Senghas, 1973) place this species in a separate genus as Tu/otis flava. The name flava means “yellow” and refers to the yellowish flowers. 38 Rhodora [Vol. 87 COMMON NAME: Tubercled Orchid FLOWERING DATES IN NEW HAMPSHIRE: late June through July DescRIPTION: A glabrous, leafy plant from several fibrous roots. Leaves 2-5, oblong-elliptic to lanceolate, spreading, becoming brac- teate above. Inflorescence sub-scapose. Floral bracts lanceolate, acuminate, longer than the lower flowers, equalling or shorter than the upper flowers. Flowers yellowish-white. Dorsal sepal ovate, concave. Lateral sepals ovate, oblique. Lateral petals ovate, forming a loose, wide hood with the dorsal sepal. Lip oblong-quadrate with a lobe-like tooth on each side near the base, and a projecting tu- bercle at the middle in front of the nectary. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). Platanthera flava is an orchid that is neither beautiful nor com- mon in New Hampshire. It is primarily a species with a more south- ern distribution, with our more robust northern. plants being referred to var. herbiola. ln New Hampshire the Tubercled Orchid grows in swales, or in wet or rocky woods. It is most often found in swales where it grows amongst a tangle of grasses, sedges, and ferns. To distinguish the orchid from the surrounding herbage takes a sharp eye, and it is helpful if one has a previous acquaintance with the Tubercled Orchid. The majority of New Hampshire stations for this species are in Strafford County, but it has also been found in Cheshire and Coos County. Platanthera flava is listed as “threatened” in Endangered and Threatened Plants of the United States (Ayensu and DeFilipps, 1978) and is presently considered “rare” in New England (Crow et al., 1981). However, the federal government has dropped it from further consideration (Federal Register , Vol. 45, No. 242, Monday, December 15, 1980). Ames and Correll (1943) have reported on the taxonomic prob- lems in this taxon that have been exacerbated by the superficial resemblance of it to Coe/oglossum viride, especially in dried mate- rials. Baldwin (1894) offered an easy way to separate the two species by stating that Platanthera flava “carries a spur, and H. viridis (C. viride] a bag.” It is true that the short sac-like spur of the latter species is a consistent character that is usually still apparent in dried plants. 1985] Brackley — Orchids of New Hampshire 39 I have found the flowers of the Tubercled Orchid to be lightly fragrant, but Morris and Eames (1929) found them “deliciously sweet-scented, with the fragrance of Madonna Lilies.” The charac- teristic tubercle, described by Gray (1862-1863) as a “nasiform pro- tuberance,” is located at the base of the lip at about the point where the lip turns sharply downward. This projection apparently func- tions to deflect pollinators so that they must approach the nectary laterally. The Tubercled Orchid is reported to be pollinated by sev- eral moths, and is also visited by mosquitoes (Leur, 1975). Offset plants may be formed vegetatively as well (Case, 1964a). GENERAL DISTRIBUTION: Nova Scotia west to Ontario, south to central Florida, west to Texas. 10. Platanthera dilatata (Pursh) Lindl. ex Beck, Bot. U.S. 347. 1833. This northern plant was described from a specimen collected in Labrador and was originally named Orchis dilatata by Pursh (1814). American botanists have traditionally treated this taxon as Habena- ria dilatata. The name dilatata is derived from Latin and means “widened,” referring to the broad base of the lip. COMMON NAMES: Bog Candle, Tall Leafy White Orchid FLOWERING DATES IN NEW HAMPSHIRE: June through August DESCRIPTION: Plant from fleshy roots, glabrous, leafy, slender or stout. Leaves several, linear to lanceolate, clasping stem below becoming bracteate above. Inflorescence a loosely or densely flow- ered cylindrical raceme. Floral bracts lanceolate, usually ascending. Flowers white, fragrant. Dorsal sepal ovate, obtuse, concave. Lat- eral sepals elliptic-lanceolate, falcate, connivent with dorsal sepal. Lip linear-lanceolate, abruptly dilated at base. Spur cylindrical, about equal in length with the lip. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). The Bog candle is a species that presents itself in a variety of forms. Sometimes it appears as a small, delicate wand with a few crystalline blossoms, while a neighboring plant from the same habi- tat may be a robust, leafy individual, with a stout raceme crowded with scores of flowers. Schrenk (1975) stated that the floral propor- 40 Rhodora [Vol. 87 tions remain constant despite the wide range of dimensions in P. dilatata, therefore the characters that distinguish it will hold true. The major difficulty in the identification of this taxon is caused by the similarity of the Bog Candle to the Tall Leafy Green Orchid, Platanthera hyperborea. In the field there is no problem as the flowers of the Bog Candle are snow-white and fragrant, whereas those of the Tall Leafy Green Orchid are green and scentless. Another character that serves to distinguish the two taxa is the shape of the lip, which in P. di/atata is typically abruptly widened at the base. In contrast, the lip in P. Ayvperborea gradually widens toward the base. The confusion between Platanthera dilatata and P. hyperborea is further amplified by the fact that the two frequently hybridize. Plants with intermediate attributes can be ascribed to P. Xmedia. In western North America many forms of the Bog Candle occur, some of which have been described as varieties. As the common name implies, this species prefers wet, peaty habi- tats. “No orchid loves better to dig its toes in the ooze and even paddle over the ankles in flood pools” (Morris and Eames, 1929). While it can be found in the southern sections of the state, it is most frequently found in Coos County (Pease, 1964), where it grows rather commonly in sloping graminoid fens and waysides. Another common habitat of this species is the damp, misty ravines of the White Mountains. Platanthera dilatata displays an interesting phenomenon in that the apex of the lip is often “caught-up” in the hood formed by the dorsal sepal and the lateral petals. As a result the flower has an odd, half-opened appearance. The strong fragrance of the flowers is usually described as “clove- like,” and is both sweet and spicy-smelling. Fernald (1926) reported that P. dilatata is locally called “Scent-bottle” in Newfoundland. Although the Bog Candle attracts “armies” of Tiger Swallowtails [ Papilio glaucus (Papilionidae)] to its blossoms (Morris and Eames, 1929), the only verified pollinators at this date are several species of moths and skippers (Leur, 1975). GENERAL DISTRIBUTION: Greenland, west through boreal North America to the Aleutians, south to New Jersey in the east, extending southward in the western mountains to California, Colorado, and New Mexico. 1985] Brackley — Orchids of New Hampshire 4] 11. Platanthera hyperborea (L.) Lindl., Gen. Sp. Orchid. 287. 1835. Linnaeus originally named this species, known to him from Ice- land, Orchis hyvperborea. Like all of our other Platantheras, it was formerly referred to the genus Hahbenaria. The epithet hyperborea means “beyond the north,” as this orchid grows within the arctic circle (Correll, 1950). COMMON NAMES: Tall Leafy Green Orchid, Tall Northern Green Orchid FLOWERING DATES IN NEW HAMPSHIRE: July through August DescRIPTION: Plant from fleshy, fusiform roots, glabrous, leafy, slender or stout. Leaves several, cauline, oblong-elliptic to linear- lanceolate, gradually reduced to bracts. Inflorescence a spike cylin- drical in shape, elongated, few- to many-flowered. Floral bracts lanceolate, usually spreading. Flowers green to greenish-yellow, not fragrant. Dorsal sepal ovate-elliptic, concave. Lateral sepals lanceo- late, oblique, spreading, reflexed. Lateral petals lanceolate, falcate, acute, erect, connivent with dorsal sepal. Lip elliptic, fleshy, gradu- ally tapering from widened base to obtuse apex. Spur cylindrical, usually shorter than lip. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 42 The Tall Leafy Green Orchid, like the Bog Candle, is a polymor- phic species. Often plants of Platanthera hyperhorea that occur in the woods are small and slender with only a few flowers, but in wet, open, roadside ditches it is often tall, thick and robust, even to the point where it has been compared to “vigorous young stalks of maize” (Case, 1964a). Some of this dimorphism is probably related to age of individual plants, as both forms may occur in the same habitat. Our plants of Platanthera hyvperborea are referable to var. huro- nensis, an overall larger and more robust plant than the typical var. hyperborea. In New Hampshire, the Tall Green Leafy Orchid occurs in wet, peaty soils, often along with P. di/atata and the hybrid P. Xmedia. There are locations where the hybrids have been observed to greatly outnumber the parent species (Schrenk, 1978). The green flowers of this orchid are scentless. It is reported that the Tall Leafy Green Orchid regularly self-fertilizes (Gray, 1862- 1863; Hagerup, 1952); however, some insects must find the flowers, 42 Rhodora [Vol. 87 attractive enough to carry pollinaria to Platanthera dilatata, or vice-versa. GENERAL DISTRIBUTION: Arctic and Boreal North America, south to Pennsylvania, west through the Great Lakes area to Washington, southward in the western mountains to California, Colorado, and Arizona; Iceland and east Asia. 5. Epipactis Zinn, Catal. 85. Mai, 1757. (conserved name) The name Fpipactis goes back to Theophrastus, who applied the name to a plant (of uncertain identity) which was used to curdle milk. There is some confusion as to the correct author of this genus but it appears that the first publication of the name after the starting point for nomenclature was in Catalogus plantarum horti academici ef agri gottingensis (Zinn, 1757). Although Zinn did not use bino- mials for the species he described, his generic names must be accepted if they are otherwise in accordance with the International Code of Botanical Nomenclature (Stafleu, 1967). The genus is comprised of about 20 species. One of these, Epipac- tis gigantea, 1S a native of western North America. The common Helleborine orchid of northeastern North America is not indigenous but an exotic, a plant introduced from Eurasia which has natural- ized here. |. Epipactis helleborine (L.) Crantz, Strip. Austr. ed. 2. Fasc. 6: 467. fig. 6. 1769. This species was called Serapias helleborine by Linnaeus, who based his description on European material. In 1769 the species was renamed by von Crantz. The specific epithet he//eborine means “like a hellebore,” referring to a group of plants in the buttercup family which the orchid supposedly resembles. COMMON NAME: Helleborine FLOWERING DATES IN NEW HAMPSHIRE: July through August DESCRIPTION: Plant leafy, from a short rhizome with many fibrous roots. Leaves clasping, spirally arranged, variable but usually elliptic-lanceolate, reduced above and changing into floral bracts. Inflorescence few- to many-flowered, a one-sided raceme. Rachis 1985] Brackley — Orchids of New Hampshire 43 lightly pubescent. Flowers small with thick, sessile ovaries sub- tended by lanceolate bracts that exceed the flowers in length. Sepals ovate-lanceolate, spreading, concave, greenish-white often with rose-purple streaks. Lateral petals ovate-elliptic, greenish-white to pink. Lip divided into two sections by a central constriction, lower half (epichile) thickened, saccate, triangular, dark purple; apical section (hypochile) thin, triangular-ovate, concave, pink. Column short, white, fleshy. Capsule obovate-ellipsoid, pendant. CHROMOSOME NUMBER: 2n = 38. Long used as a medicinal herb in Eurasia, Epipactis helleborine was best known in the Old World as a remedy for gout. The Helle- borine is the only widespread non-native orchid in our flora.* How it first immigrated to this continent is a mystery, but when Hellebo- rine was discovered in 1879, it had already become established in a deciduous woods near Syracuse, N.Y. (Anon., 1879). Mrs. M. P. Church of the Syracuse Botanical Club surprised the professional botanists of her day by making this remarkable find. Three years later, another amateur botanist recorded a well-established colony of Helleborine near Scajaquada Creek, Buffalo (Zenkert, 1950). In subsequent years populations of Helleborine were reported from Toronto and Montreal (Leur, 1975). Epipactis helleborine was clearly here to stay. In aggressive, weed-like fashion, Helleborine continued to extend its range. One state after another—Massachusetts, 1902; Indiana, 1931; Missouri, 1940—added the “new” orchid to their state lists (Doyon and Cayouette, 1966). Helleborine was found in Hartland, Vermont near the New Hampshire state line in 1925, but was not recorded in this state until 17 years later. In 1942, in Plainfield (Sullivan County), directly across the river from Hartland, Vermont a single plant of Helle- borine growing on a shaded bank provided the first record of its occurrence in New Hampshire (Upham, 1942). In the Jessup Herbarium at Dartmouth College in Hanover, N. H., there is a map of the state studded with colored pins which represent *The recent reports from Canada (Elliott and Cook, 1970; Anderson and Goltz, 1982) of the occurrence of the European orchid Listera ovata indicate that a similar invasion may be taking place by that species. 44 Rhodora [Vol. 87 records of the occurrence of E. helleborine; each color designates an interval of time since 1942.'Dr. James Poole, Professor Emeritus of Botany at that institution, has provided a graphic illustration of the procession of this orchidaceous weed through time within the state. Today, Helleborine can be found in every county in New Hampshire. Helleborine is reported to prefer circumneutral soils (Correll, 1950), but Stucky (1967) indicated that the presence of free carbon- ates in the substrate is a more important ecological requirement. | have found, as she did, Helleborine growing amongst old founda- tions, but the most likely place to encounter this orchid in New Hampshire is in rich, deciduous woods. In testimony to its aggres- siveness, Helleborine is able to flourish in a wide variety of habitats. The most robust specimens I ever observed were waist-high, grow- ing in open pine woods in Belknap County. Immature Helleborine plants produce only a leafy stalk which somewhat resembles that of Cypripedium arietinum. Each year the plants become larger until they become reproductive (Leur, 1975). A mature plant produces many, rather inconspicuous, small flowers. Epipactis helleborine produces large amounts of nectar in the cup-like hypochile (L. A. Nilsson, 1978). The lower half of the lip, the epichile, provides a platform for insects to perch on while par- taking of the nectar. As the insect (in this case a wasp) backs out of the flower, its head comes in contact with the sticky rostellum. The rostellum wall ruptures and its sticky effluent glues the pollinia to the insect’s head (Darwin, 1877). In Europe, Darwin (1877) observed the wasp Vespa sylvestris (Vespidae) pollinate Epipactis helleborine. On this continent wasps of the genus Vespula (Vespidae) have been reported as the most important pollinators (Judd, 1972; Leur, 1975). Hagerup (1952) has found that populations of Helleborine in Denmark are autogamous, and that this process takes place directly before or just after the flower opens. Helleborine produces abundant ripe capsules filled with many microscopic seeds. The fact that the seeds are easily dispersed far and wide is documented by the rapid spread in northeastern North America of this most interesting and delightful orchid. GENERAL DISTRIBUTION: Eurasia and northeastern North America, west to Michigan and Missouri. 1985] Brackley — Orchids of New Hampshire 45 6. Listera R. Brown in W. et W. T. Aiton, Hort. Kew. ed. 2, 5: 201. 1813. (conserved name) This genus of minute orchids was named in honor of Martin Lister, a 17th century English naturalist and physician. Although the genus is widely distributed in temperate regions, most of the approximatley 25 species are rare and local. Twayblades are extremely small and have tiny, inconspicuous flowers. All the Listeras that occur in the northeast have an elon- gated somewhat bifurcated lip which secretes nectar along a central groove. While some Listeras are apparently scentless, most of our species produce fetid odors that have been described as “truly repul- sive” (Ramsey, 1950; Ackerman and Mesler, 1979). Pollination biology of the Twayblades is especially fascinating as it features an active attachment of the pollinia to the pollinator. This process begins in the bud when the anther opens and drops the two pollinia onto the concave back of the rostellum (Baldwin, 1894). The rostellum, which forms a hood-like cover over the stigmatic surface, has tiny projections on the front margin that act as triggers (Ramsey, 1950). When the triggers are touched, even lightly, the rostellum explosively expels a viscid, quick-drying fluid onto “the object whose touch caused the exudation” (Miller, 1883). Instantly, the margins of the rostellum fold back releasing the pollinia which fall precisely onto the drop of quick-drying cement (Ackerman and Mesler, 1979). When this mechanism acts on a pollinator, it frightens the insect which quickly flies away bearing a crown of pollinia. In their study of Listera cordata, Ackerman and Mesler (1979) found that after the pollinia are removed, the rostellum spreads flat over the stigma for approximately one day. This position of the rostellum assures that the flower does not become self-pollinated by a returning insect. Gradually the rostellum rises to expose the now- receptive stigmatic surface. Cross-pollination occurs when an insect bearing pollinia from another Twayblade alights to partake of the nectar and brings a pollinium in contact with the stigma. Since only small pieces of the pollen-masses adhere to the sticky stigma, it is possible for several flowers to be pollinated by a single insect (Ack- erman and Mesler, 1979). This precise pollination mechanism of Listera is adapted for use with unspecialized pollinators (Proctor, 46 Rhodora [Vol. 87 1978). European Twayblades are pollinated by ichneuman wasps, beetles, and small flies (Summerhayes, 1951). In our flora, there are three species of Listera. In addition, a hybrid between L. convallarioides and L. auriculata has been recently described as L. Xve/tmanii (Case, 1964b). Cody and Monro (1980) reported this hybrid as occurring in Coos County, N.H. Key to the Species of Listera in New Hampshire 1. Lip linear-oblong with a pair of horn-like teeth at the base, divided at least half-way to base into 2 linear prongs; leaves VANE COT OLE a oy dk Gees 445d eR eee OR 1. L. cordata 1. Lip oblong to obovate without horn-like teeth, cleft less than half-way to base into oblong or broadly rounded lobes; leaves ovate-elliptic to suborbicular........... 0.0 c cece ee eees (2) 2. Lip oblong, broadest at base with auricles clasping the column; pedicels. glaQrous cs cise teess veed seas 3. L. auriculata 2. Lip obovate, broadest at apex with a slender claw at base and a triangular tooth on each side near claw: pedicels BIANOUIOE so axcdandne bien neve ekeas 2. L. convallarioides 1. Listera cordata(L.) R. Brown in W. et W. T. Aiton, Hort. Kew., ed. 2, 5: 201. 1813. The Heart-leaved Twayblade was described by Linnaeus based upon European plants of cold, moist forests. He placed this species (and another European Twayblade, L. ovata) into the genus Oph- rvs. Robert Brown realized that the Twayblades were distinct from Ophrys and separated them to the genus Listera. As the common name indicates, the specific epithet means “heart-shaped.” COMMON NAME: Heart-leaved Twaybade FLOWERING DATES IN NEW HAMPSHIRE: June through July DESCRIPTION: Plant slight, 10-20 cm tall, mostly glabrous. Roots few, fibrous. Leaves two, opposite, ovate-cordate, midway up stem. Inflorescence a slender raceme. Flowers inconspicuous, greenish to reddish-purple, subtended by minute bracts. Sepals and lateral petals ovate-oblong to elliptic, spreading, 2-3 mm long. Lip linear with two horn-like teeth at base, deeply cleft for half of its length into two spreading, lanceolate prongs. Column small, stubby, 0.5 mm long. Capsule ovate, ribbed, semi-erect. 1985] Brackley — Orchids of New Hampshire 47 CHROMOSOME NUMBERS: 2n = 38, 42. The circumboreal distribution of Listera cordata makes it the most widespread species of the genus. A diminutive inhabitant of wet, mossy Spruce-Fir woods and Cedar swamps, the Heart-leaved Twayblade is easily overlooked by the orchid hunter. The plant is often buried so deeply in loose moss that the twin heart-shaped leaves appear to be basally located. The minute flowers are spread widely to expose the nubby column and the curious pair of horn-like teeth at the base of the lip. The microscopic dimensions of the Heart-leaved Twayblade and its production of a fetid odor usually prevents this species from being sought out as an addition to wild- flower gardens. Ackerman and Mesler (1979) stated that the flowers of Lisrera cordata are self-compatible but not autogamous. In their study of northern Californian populations of the Heart-leaved Twayblade, they found the most common pollinator to be fungus gnats of the genera Mycetophila (Mycetophilideae), Sciara and Corynoptera (Sciaridae). That these small diptera can serve as effective pollina- tors, was illustrated by the high percentage of fruit set in L. cordata, especially when contrasted with percentage fruit set for other allog- amous orchids (Mesler ef a/., 1980). The Heart-leaved Twayblade is a prolific seed producer. One plant can yield as many as 2,860 seeds a year (Stoutamire, 1964). Vegetative reproduction may also occur in this species (Case, 1964a). In New Hampshire, L. cordata has beeti most often collected in the White Mountains and northern Coos County. There have been few modern collections of the Heart-leaved Twayblade in the state. Two of the most recent records are from the northernmost town of Pittsburg, collected by the late Albion R. Hodgdon of the Univer- sity of New Hampshire. Storks and Crow (1978) listed Listera cordata as “widespread but infrequent.” It is possible that in some areas of the White Mountains where hiking (and botanizing) activities are high, some populations of the Heart-leaved Twayblade may have been diminished, but other populations of L. cordata, with their delicate stems and tiny flowers, surely grow unnoticed in scattered locations within the state. 48 Rhodora [Vol. 87 GENERAL DISTRIBUTION: Circumboreal; in North America extend- ing south along the mountains to North Carolina, California, and Colorado. 2. Listera convallarioides (Sw.) Torrey, Comp. Fl. North and Mid- dle States. 320. 1826. A North American specimen was described by Swartz (1800) as Epipactis convallarioides. In 1818, Nuttall simply listed L. convalla- rioides as the third species in the genus Listera without a description or reference to the original author and description. Because of Nut- tall’s lack of description, his name must be regarded as invalid, according to Article 33 of the /nternational Code of Botanical Nomenclature (Stafleu et al., 1978). Therefore, Torrey, who de- scribed Listera convallarioides in 1826, can technically be consi- dered the author of this species, but the matter needs further study. Torrey’s use of the epithet convallarioides indicates that he was familiar with Swartz’s taxon, and may have intended to transfer it to Listera. The epithet convallarioides means “like a Convallaria,” re- ferring to a supposed resemblance to Lily-of-the-Valley. COMMON NAMES: Broad-leaved Twayblade, Lily-leaved Twayblade FLOWERING DATES IN NEW HAMPSHIRE: July DESCRIPTION: Plant usually slender, 5-30 cm tall (averaging 16 cm tall here), glabrous below leaves, pubescent above. Leaves 2, oppo- site, about half-way up stem, elliptic to sub-orbicular. Inflorescence a loose, many-flowered raceme. Floral bracts lanceolate, semitrans- lucent. Flowers yellowish-green, translucent, on slender glandular pedicels. Sepals linear-lanceolate, reflexed. Lateral petals narrower than the sepals, linear, reflexed. Lip conspicuous, much elongated, wedge-shaped with a central groove, widely flared and bilobed with a minute tooth in the sinus, auriculate near the base and narrowing into a slender claw. Margins of lip minutely ciliate. Column recurved, winged. Capsule ellipsoid. CHROMOSOME NUMBER: 2n = 36 (Moore, 1973). The Broad-leaved Twayblade is a northern plant of cold soils (Case, 1964a). In New Hampshire this species inhabits mossy woods and Cedar (Thuja occidentalis) swamps. The shiny, broad green leaves appear early in the summer well before the flowering spike 1985] Brackley — Orchids of New Hampshire 49 elongates. The unusual-looking flowers have inconspicuous sepals and lateral petals that are reflexed from the oversized bilobed lip and overhanging column. The blossom is stiffly perched on the thread-like pedicel with the lip held at a 45 degree angle from the stem. Unfortunately, I have found no specific references to insect vis- itors of Listera convallarioides and have not observed pollination in this species myself. An interesting observation on the inflorescence of the Broad-leaved Twayblade is that it strongly resembles a raceme of small diptera with their wings held in a resting position. Comparing a dorsal view of a fungus gnat (Mycetophilideae) with the lip of L. convallarioides, similarities of color, translucence and the ciliated margins are apparent. The dark central line of the lip corresponds to the very narrow body of the insect. A detailed study of the pollination biology of this species might reveal some interest- ing relationships with insect visitors. All locations for the Broad-leaved Twayblade in New Hampshire are within Coos County. There have been very few collections of this species in the state in the past 50 years. Case (1964a) reported finding many individuals of Listera convallarioides in the western Great Lakes area with blackened stems, and Wherry stated that this species is “extraordinarily susceptible to attack by parasitic fungi” (Correll, 1950). GENERAL DISTRIBUTION: Boreal North America; mostly Canada, reaching northern New England and the northern Great Lakes states. Disjunct to the mountains of North Carolina and Tennessee. In the west; north to Alaska, south to the mountains of Utah and Arizona. Closely related to L. makinoana of Japan (Leur, 1975). 3. Listera auriculata Wiegand. Bull. Torr. Bot. Club 26: 166, pl. 356, fig. 2. 1899. Wiegand described L. auriculata from material collected in Quebec, New Hampshire, and Maine. The epithet auriculata means “ear-like,” in reference to ear-like appendages which curl around the column. COMMON NAME: Auricled Twayblade FLOWERING DATES IN NEW HAMPSHIRE: Mid-June to mid-July 50 Rhodora [Vol. 87 DESCRIPTION: Plant slight, slender. Stem glabrous below leaves, pubescent above. Leaves 2, opposite, ovate-elliptic, dark green, sit- uated above the middle of the stem. Inflorescence a loosely flowered raceme. Floral bracts small, approximately 5 mm, oblong-lanceolate. Flowers watery blue-green on stout, glabrous pedicels. Sepals and lateral petals reflexed from column and lip. Dorsal sepal elliptic- obovate, blunt at apex. Lateral sepals lanceolate-elliptic. Lateral petals linear, obtuse. Lip oblong, somewhat narrowed in the center, ciliate at margins, cleft at apex into two blunt lobes with a narrow sinus. Base of lip forming two auricles which curve behind the column. Column stout, curved toward lip. Capsule ellipsoid. CHROMOSOME NUMBER: 2n = 36. Listera auriculata is a rare orchid with a distribution restricted to northeastern North America. It is quite close in appearance to L. borealis, the Northern Twayblade, which occurs in Canada and western North America. Where the two species grow together they can be distinguished by the shape and position of the auricles. In L. auriculata, the rounded auricles curve behind the column while in L. borealis the oblong auricles diverge from the column. The Auricled Twayblade has a distinctive habitat preference for alluvial thickets where it grows under Alder (A/nus viridis var. crispa) shrubs. In such a thicket, I discovered a population of Auricled Twayblades pushing through leaf litter that had accumu- lated on the sandy soil. The largest plant was less than 10 cm tall, yet their pale blue-green flowers were beginning to unfurl. This riverside site is only the fifth station and the only known extant site in New Hampshire for this rare, small orchid. All recorded locations in the state for this species occur within Coos County. Northern sections of Maine, New Hampshire and Vermont form the southeastern limit of the range of this boreal species. The scar- city of the Auricled Twayblade has earned it a listing as “Endan- gered” for New England (Crow et al., 1981). Ona national scale, L. auriculata is currently under review for listing as an Endangered or Threatened Species of the United States (Federal Register, Vol. 45, No. 242, Monday, Dec. 15, 1980). GENERAL DISTRIBUTION: Northeastern North America; Newfound- land south to northern New England, west to Michigan. 1985] Brackley — Orchids of New Hampshire 51 7. Triphora Nuttall, Gen. N. Am. Pl. 2: 192. 1818. Once included within the genus Pogonia, this genus of small woodland orchids was segregated in 1818 by Nuttall. The genus occurs exclusively in the Western Hemisphere and includes about 10 species. Case (1964a) stated that “structurally and physiologically members of this genus appear to be intermediate between independ- ent, free-living plants and dependent saprophytes.” The name 7ri- Phora is from two Greek words meaning “three-bearing,” referring to the number of flowers per plant in bloom at one time. 1. Triphora trianthophora (Sw.) Rydb. in Britton, Man. 298. 1901. This North American species was first described in 1800 by Swartz as Arethusa trianthophora. The Three-birds Orchid has also been known as Pogonia pendula which translates as “Nodding Pogonia,” another common name for this species. The specific epithet trianthophora is somewhat repetitive as it means “bearing three flowers.” COMMON NAMES: Three-birds Orchid, Nodding Pogonia FLOWERING DATES IN NEW HAMPSHIRE: mid-August to early September. DESCRIPTION: Plant stoloniferous, producing white ovoid tubers and slender white roots. Stem glabrous, greenish-purple, succulent. Leaves alternate, ovate, clasping stem. Inflorescence a raceme of 1-6 pedicellate flowers borne in the axils of the upper leaves. Flowers fragile, spreading, white or pinkish-white. Dorsal sepal oblanceo- late. Lateral sepals and lateral petals oblanceolate, falcate. Lip with a slender claw, three-lobed. Lateral lobes ovate, obtuse. Mid-lobe ovate with sinuate margins. Column white with terminal anther. Capsule ellipsoid, erect. CHROMOSOME NUMBER: 2n = 18 (Baldwin and Speese, 1957). The Three-birds Orchid is so named for the three flowers on each plant that are usually in bloom at the same time—“one perfect bloom with a fading flower below and an opening bud above” (Morris and Eames, 1929). The blossoms of the tiny, fragile orchid range in color from waxy white to rosy pink. Instead of the usual yellow-colored pollen, the Three-birds Orchid is unique in having 52 Rhodora [Vol. 87 pollen masses that are violet. In keeping with the scheme of threes, the lip has three lobes and a median trio of long green crests. The only member of the genus that occurs north of Florida, 7ri- phora trianthophora is widespread but uncommon throughout its range (Porcher, 1977). In addition to being uncommon, sizes of known populations of the Three-birds Orchid fluctuate widely from year to year. The plant may be abundant at a particular station during one year and be absent or scanty for the next few years (Eastman, 1976). An example of the varying size of Triphora trian- thophora populations is furnished by one well-known New Hamp- shire station where 5000 plants were found in 1916 but only 5 of the orchids were located in 1978 (Lowenstein, 1917; Storks and Crow, 1978). At the same location in 1980 I counted approximately 200 plants in flower. The reason for the elusive occurrence of flowering plants of Tri- phora is attributed to the largely subterranean existence of this orchid (Ames, 1922a). During most of its life, this species reportedly remains underground ina tuberous condition. In this state the plant exhibits a saprophytic mode of existence, deriving its nourishment from the mycorrizal association present in the tissues of the tuber. Because colonies of the Three-birds Orchid also produce green stems and leaves capable of photosynthesis during its brief life above ground, the plants have been termed “hemisaprophytes” or “semisaprophytes” (Batchelder, 1909; Case, 1964a). Triphora trianthophora requires a specialized habitat in northern New England. It is often found in nearly pure stands of Beech (Fagus), where it grows in hollows and depressions that have filled up with fallen leaves. Prince (1902) described the Three-birds Orchid at Intervale (Conway) New Hampshire as “often pushing up through the Beech leaves and carrying them like an unwieldly collar at a little distance below the flowers.” The companion herbs of this species are also largely saprophytic and parasitic. Indian Pipes (Monotropa uniflora), Spotted Coral-root (Corallorhiza maculata) and Beech-drops (Epifagus virginiana) commonly share the same Beech grove. In late August Triphora trianthophora begins to make its tran- sient appearance above ground. The plants grow rapidly for about two weeks and quickly develop buds. A drop in the average night temperature may serve as a stimulus to flowering, as 48 hours later 1985] Brackley — Orchids of New Hampshire = all of the mature buds in a colony will simultaneously open (Leur, 1975). Leur also stated that the strategy of mass flowering increases the chance for the (literally) ephemeral flowers to be cross- pollinated. Lowes (1920) listed the small bee Halictus quadrimacu- latus (Halictidae) as a pollinator of the Three-birds Orchid in New Hampshire. A number of seed capsules are commonly observed after the flowering season. Triphora trianthophora also reproduces vegetatively by the production of stolons that develop new tubers (Ames, 1922a). In New Hampshire, the Three-birds Orchid is listed by Storks and Crow (1978) as “widespread but infrequent.” Although it has been occasionally found in scattered locations throughout the state, most populations of this orchid occur within Carroll County. GENERAL DISTRIBUTION: New England and southern Ontario, west to Michigan and Wisconsin, south to Florida, west to Texas. 8. Goodyera R. Br. in W. et W. T. Aiton, Hort. Kew. ed. 2, 5: 197. 1813. (conserved name). This genus was proposed by Robert Brown who named it in honor of the botanist John Goodyer (1592-1664) who “englished” Dioscorides’ herbal (Freeman, 1943). There are about 25 species of Goodyvera widely distributed in temperate and tropical regions of the world (Correll, 1950). Members of this genus are characterized by creeping rhizomes and rosettes of evergreen leaves. The leaves are similar in shape, though much smaller, to the Common Plantain (Plantago) and are marked with reticulations reminiscent of the patterns of snakeskins, hence the common name of Rattlesnake Plantains. Of the four North American species, three are found in New Hampshire. The Rattlesnake Plantains are capable of hybridizing and appar- ently do so quite often in the Great Lakes region (Case, 1964a). Because of slightly different habitat preferences and less geographi- cal overlap this phenomenon is less likely to occur in our area. Key to the species of Goodyera in New Hampshire I, Raceme one-sided or loosely spiralling iv iaccxcwaevevcawa (2) 2. Leaves less than 3 cm long; raceme one-sided............. OT eee ee re 1. Goodyera repens var. ophioides 54 Rhodora [Vol. 87 2. Leaves 2-7 cm long; raceme loosely spiralling ............ Terre Tere rr eT er Tree Tee 2. Goodyera tesselata 1. Raceme densely flowered, cylindrical in overall appearance... Ad ES REARS EV ULAR EEE SRERE EL OES RAI EE ENR EE EIS NE (3) 3. Beak of rostellum equal in length or greater than body of stigma; leaves bluish-green ........ 2. Goodvera tesselata 3. Rostellum essentially beakless; leaves dark green.......... Se aeNN Eee Ke eeae ead os © Oe OORVERE PUDESCENS 1. Goodvera repens (L.) R. Br. in W. et W. T. Aiton, Hort. Kew. ed. 2. 5: 198. 1813. Described by Linnaeus as Satyvrium repens trom plants collected in Sweden and Siberia, this species is widespread in the cool moist coniferous forests of Eurasia. The typical var. repens with plain green leaves is also found in western North America. Our eastern North American plants are referable to var. ophioides Fern. This variety (Fernald, 1899) is distinguished from var. repens by its smaller size and conspicuous white markings that border the veins of the leaf. The epithet repens translates as “creeping,” descriptive of the growth habit of the rhizome. In Europe this species is known as “Creeping Ladies’-tresses” (Summerhays, 1951). The name ophi- vides means “snake-like,” alluding to the patterns on the leaves which “strongly suggest the markings of a serpent” (Fernald, 1899). COMMON NAMES: Lesser Rattlesnake Plantain, Creeping Rattle- snake Plantain FLOWERING DATES IN NEW HAMPSHIRE: mid-July through August. DeEscCRIPTION: Plant from a creeping rhizome, scapose, rosette- forming, approximately 16 cm tall. Leaves 3-7 on short pedicels, ovate to oblong-elliptic with white reticulations. Inflorescence a one-sided raceme, loosely flowered. Floral bracts lanceolate, acu- minate. Flowers white to geenish-white, pubescent on outer surfa- ces. Dorsal sepal ovate, concave. Lateral sepals oblique. Lateral petals oblong-spatulate, connivent with dorsal sepal and forming a hood over lip. Lip globose-saccate with flaring margins and an elongated recurved tip. Column with beak shorter than body of stigma. Capsule ovoid. CHROMOSOME NUMBER: 2n = 30. 1985] Brackley — Orchids of New Hampshire 55 This orchid is found in dark coniferous woods throughout its range. The creeping manner of growth in this and in all Goodyera species derives from horizontally-growing rhizomes which are located just below the surface of the forest floor. Upright plants develop at the tips of rhizomes. According to Summerhays (1951) the production of a rosette of green leaves takes five years from the time of germination, and it is only after eight years that a flowering stalk appears. Other workers believe that there may be a shorter time requirement for the initiation of flowering (Case, 1964a). Vegetative plants of this species are more commonly observed than flowering individuals. Downie (1940) found that plants of G. repens remain dependent on mycorrhizae for nourishment throughout their lives. The slender, one-sided flower stalk has up to 20 small sweet- smelling blossoms subtended by bracts. A close look at one of the white pubescent flowers reveals a lip in the form of a boat-like pouch. The apex of the lip extends out into a pointed channel, much like the pouring groove of a pitcher. Nectar is produced inside this vessel, an apparent adaptation for insect pollination, Goodyera species are protandrous, a reproductive syndrome which promotes cross-fertilization. In the newly-opened blossoms of the Lesser Rattlesnake Plantain the column is bent forward, cover- ing the stigma and almost touching the lip. This arrangement pro- vides an opening only large enough for a bee’s proboscis. A bee stopping to sip nectar will immediately come in contact with a viscid gland which glues the pollen-masses to the bee’s mouth (Gibson, 1905). As the blossoms age the column turns upward and backward causing the flower to open wider and expose the newly receptive stigma (Gray, 1862-1863). Any pollinia-bearing bee will now come in direct contact with the stigma and effect cross-pollination. This same process apparently occurs in the other North American members of this genus (Ackerman, 1975). Bumblebees (Apidae) are responsible for fertilizing the Lesser Rattlesnake Plantain. Darwin (1877) observed Bombus pratorum with pollinia from Goodyera repens, and Miller (1883) saw this orchid being visited by B. mas- trucatus in the Alps. There are also reports of Bombus visiting North American plants of the Lesser Rattlesnake Plantain (Acker- man, 1975). 56 Rhodora [Vol. 87 Because of the relatively high percentage of seed set in Goodyera repens, it has been suggested by Hagerup (1952) that, in Denmark, this species may be autogamous. Bagging experiments on G. repens var. ophioides by Kallunki (1976) in northern Michigan indicated that this is not the case in eastern North American plants. The Lesser Rattlesnake Plantain is a successful plant from the stand- point of reproduction with a high (for orchids) percentage seed-set and the additional ability to propagate vegetatively by its horizontal rhizomes. Of our Goodyeras, Goodyvera repens is by far the most widely distributed. Variety repens is found throughout Eurasia and in western North America. Our var. ophioides very rarely occurs in Europe but is known from Asia, especially in the Himalayan Moun- tains in India (Correll, 1950). On this continent var. ophioides grows in the northeast and in Alaska. Kallunki (1976) proposed that the taxon survived glaciation in refugia near the glacial edge, perhaps in the Appalachian Mountains. The Alaskan plants may well have been an element of the boreal forest flora that survived glaciation in the ice-free area connecting Alaska and Siberia, known as Beringia (Murray, 1980). In New Hampshire the Lesser Rattlesnake Plantain favors the cool forests of the White Mountains and northern New Hampshire. Goodyera repens var. ophioides can grow in dark moist woods where its tolerance of such conditions often results in its being one of the only herbs present in the community. GENERAL DISTRIBUTION: In the formerly glaciated areas of eastern North America, south along the Appalachians to North Carolina and Tennessee, disjunct to Alaska. Intermediate forms between the two varieties occasionally found in the western mountains (Kal- lunki, 1976). Rarely from Europe, occasionally from Asia. Very close to G. japonica from Japan (Hultén, 1962). 2. Goodyera tesselata Lodd. Bot. Cab. 10: pl. 952. Apr. 1825. Goodyera tesselata was first described by Loddiges, who pub- lished a colored figure of the plant in his Botanical Cabinet (1824) that was drawn from plants found in New York and Philadelphia. However, Loddiges’ species remained unknown to most botanists until it was included in the seventh edition of Gray's Manual of Botany (Robinson and Fernald, 1908). Northeastern plants of G. 1985] Brackley — Orchids of New Hampshire 57 tesselata collected prior to this publication were labeled either G. menziesii (presently known as G. oblongifolia) or G. repens (Fer- nald, 1899). Some of these old designations still remain on herba- rium sheets, causing much confusion about the true distribution of G. oblongifolia; this species does not occur in New Hampshire. The epithet resse/ata means “checkered,” describing the pattern on the leaves in this species. COMMON NAMES: Tesselated Rattlesnake Plantain, Checkered Rat- tlesnake Plantain FLOWERING DATES IN NEW HAMPSHIRE: — late July to early September. DESCRIPTION: Plant erect, up to 30 cm tall, densely glandular- pubescent, from a creeping rhizome with fibrous roots. Leaves with short broad petioles, in a basal rosette, light bluish-green reticulated with white. Inflorescence a loosely or densely flowered raceme, somewhat one-sided, loosely spiralled or (occasionally) cylindrical. Floral bracts lanceolate. Flowers with stout ovaries, white to greenish-white, pubescent on outer surfaces. Sepals oblong- lanceolate, oblique. Lateral petals spatulate, oblique, connivent with dorsal sepal to form a hood. Lip saccate with recurved beak. Column with slender beak equal in length or longer than body of stigma. Capsule ovoid. CHROMOSOME NUMBER: 2n = 60 (Kallunki, 1976). The Tesselated Rattlesnake Plantain is the most common Good- vera in northern New England. It was suggested by Fuller (1933) and Case (1964a) that G. resselata may have originated through post-glacial hybridization of G. oblongifolia and G. repens var. ophioides. Kallunki (1976) researched this hypothesis for her thesis and the results of her study supported the theory of a past hybrid origin for G. tesselata. Her research showed the Tesselated Rattle- snake Plantain to be intermediate in morphological characters between the two putative parents and to share their respective phe- nolic compounds. Goodyvera tesselata, with a diploid chromosome number of 60, is reported to be an allotetraploid derived from the diploids G. repens var. ophioides and G. oblongifolia (both 2n = 30), and that morphologically intermediate triploids exist (Kallunki, 1976). The presence of these triploid plants in the area of the Great Lakes states are the cause of much difficulty in identification. 58 Rhodora [Vol. 87 The establishment and proliferation of the Tesselated Rattlesnake Plantain in regions where the parental species are scanty or absent (i.e., New Hampshire) indicates that Goodyera tesselata is not just a hybrid swarm, but is clearly a distinct species (Case, 1964a). This species is larger and stouter than the Lesser Rattlesnake Plantain and is better adapted to drier situations. Instead of grow- ing in colonies, as is the habit of other Goodyeras, plants of Gooa- vera tesselata are most apt to grow singly and at some distance from each other. The late summer orchid hunter is almost certain to be rewarded by the presence of this species. This species is common in mixed or coniferous woods, especially when few other herbs are present. The narrow, white-flowered stalks are quite noticeable in shady woods. Dried flower stalks from the previous year are often found next to fresh rosettes of the watery blue-green leaves. In contrast, the flow- ering plants that I have observed never have an old inflorescence in evidence. From these observations it appears that at least one year elapses between production of a new rosette of leaves and the onset of flowering. The pollination mechanism in the Tesselated Rattlesnake Plan- tain is probably much the same as in Goodyera repens var. ophioides. Bagging experiments on this species done by Kallunki (1976) indicated that G. resse/ata does not normally self-pollinate. In the same study, bees of the genus Bombus (Apidae) were observed visiting the fragrant flowers. GENERAL DISTRIBUTION: Newfoundland west to Ontario and Min- nesota, south through New York. 3. Goodvera pubescens (Willd.) R. Br. in W. et W. T. Aiton, Hort. Kew. ed. 2. 5: 198. 1813. Willdenow (1805) first described this species as Neottia pubes- cens, and it has also been known as Epipactis pubescens. The epithet pubescens stands for “downy” describing the densely pubescent flowering stalk. COMMON NAME: Downy Rattlesnake Plantain FLOWERING DATES IN NEW HAMPSHIRE: late July through August. 1985] Brackley — Orchids of New Hampshire 59 DESCRIPTION: Plant from a creeping rhizome with fibrous roots, densely pubescent, erect, up to 37 cm tall. Leaves with a short, broad petiole, dark green with many white reticulations, ovate- lanceolate. Inflorescence a stout, cylindrical, densely flowered raceme. Floral bracts lanceolate. Flowers globose, white, with stout ovaries. Sepals ovate, concave, narrowed into a blunt apex. Lateral petals oblong-spatulate, oblique, connivent with the dorsal sepal to form a hood. Lip scrotiform with a short recurved beak. Column short, rostellum essentially beakless. Capsule globose. CHROMOSOME NUMBER: 2n = 26 (Kallunki, 1981). This species, with its dark green leaves embossed with delicate white markings, has the most handsome foliage of all our native orchids. Because of these attractive leaves, plants are often taken for terraria. Goodyera pubescens is Our counterpart of the southeast Asian Jewel Orchids (Macodes spp.) that are cultivated for their foliage. All of our Rattlesnake Plantains are termed “evergreen” but only in this species are the leaves a rich green color; our other two species have pale bluish-green leaves. The type location of this species was listed by Willdenow (1805) as “from Canada to Florida,” but the range of Goodyera pubescens only extends south to the mountains of Tennessee. This species Is quite common in New Hampshire where it grows in mixed woods. Often a colony of the Downy Rattlesnake Plantain will be found nestled at the bottom of a large old Pine tree. Goodyera pubescens grows in colonies so dense that the leaves of individual plants often overlap. Ames (1922b) investigated this habit of growth and stated that the compactness of the colony derives in part from the constant replenishment of seedlings. He concluded that only those seeds that fall within an established colony will be assured of successful germi- nation owing to the already established presence of mycorrhiza) fungi. This species also perpetuates itself vegetatively and colonies of G. pubescens usually are composed of plants in different stages of maturity with only a few flowering shoots per colony. During one season Baldwin (1894) counted 572 plants of G. pubescens, of which 103 flowered. The Downy Rattlesnake Plantain has long been used as a medici- nal plant, as evidenced by one of its old common names “Scrofula 60 Rhodora (Vol. 87 Weed.” Baldwin (1894) quoted the American Herbal published in Walpole, New Hampshire in 1801: Country people use a decoction of the leaves for skin diseases, and Captain Carver says the Indians are so convinced of its power as an antidote that they allow a snake to drive its fangs into them, then chew the leaves and apply them to the wound. It is fortunate that poisonous snakes have only been rarely reported in New Hampshire. Pursh (1814) reported that, “This plant has lately made a great noise among the country people, as infallibly curing the bite of a mad dog.” Whigham and McWethy (1980) reported that Goodyera pubes- cens is not self-pollinating and that cross-pollination is effected by Bombus spp. (Apidae). It appears that all of our Goodyera species have evolved reproductive strategies involving bumblebees of the genus Bombus. The Downy Rattlesnake Plantain produces a rela- tively high percentage of seeds, and, as previously stated, can also propagate vegetatively. Ames (1938) noted that in this species the ovaries of the fading flowers tend to untwist returning the lip to a more upright position, GENERAL DISTRIBUTION: Quebec and Ontario south through the Appalachian uplands to Georgia and Alabama, west through the Great Lakes states to Minnesota. 9. Spiranthes L. C. M. Rich., De Orch. Europe. Annot. 28. 1817. (conserved name) A large genus of about 300 species, the majority of which are confined to the New World, Spiranthes is closely allied with the morphologically similar Goodyera, although the Ladies’-tresses lack the evergreen leaves and saccate lips which characterize the Rattles- nake Plantains (Case, 1964a). In most species of Spiranthes the leaves begin to wither at, or before, flowering time. In old botany books, this genus is referred to as Gyrostachy’s or /bidium, two generic names that pre-date the name Spiranthes. The name Spiranthes is conserved over the other earlier names accord- ing to the /nternational Code of Botanical Nomenclature (Stafleu et al., 1978). 1985] Brackley — Orchids of New Hampshire 61 The species of Spiranthes that occur in our geographic area have more or less basal leaves and a spike of small flowers. All of the northeastern species have white or whitish flowers. The name Spiranthes is from two Greek words, and means “coiled flower,” referring to spiralling of the flower spike which occurs in most species. If the inflorescence appears to be composed of one vertical row of flowers spiralled around the stem, the floral arrangement is termed “one-ranked” or “single-ranked.” Two verti- cal rows of spiralling flowers are said to be “two-ranked,” and so forth. The spirals may be “right-handed” or “left-handed.” Correll (1950) found the majority of single-ranked inflorescences he exam- ined to be “right-handed.” In this genus the dorsal sepal and lateral petals usually overlap to form a hood over the lip. The base of the lip has two small round knobs called “callosities” or “tuberosities,” which are sometimes diagnostic in determining a taxon. The majority of our species are protandrous. While the flower is still in bud, the anther, which ts pressed over the rostellum, matures and releases the two pollen-masses. The pollinia fall into a boat- shaped disk that is embedded in a furrow on the rostellum (Gibson, 1905). The pollination biology of Spiranthes was studied by Darwin (1877) who found that when the furrow is touched it instantly splits open and exudes a milky adhesive. Ina matter of seconds the adhe- sive dries, cementing the pollen-bearing disk to the bee’s proboscis. The column in newly opened flowers is pressed down against the lip, blocking the passage-way to the immature stigma and nectary. A probing insect coming in contact with the rostellum may collect several pollen-bearing disks from’ different flowers before landing ona wide-open, older flower with a sticky, receptive stigma. Darwin (1877) also observed that the insects first visited the lowermost, most mature flowers on a Spiranthes spike and worked up to the uppermost, younger blossoms. On the higher blossoms the bees encounter the pollinia and carry them off to the wide-open lower- most flowers of another plant, thus effecting cross-pollination. The pollinators of most Spiranthes species are reported to be bumble- bees (Darwin, 1877; Ames, 1921). Spiranthes lucida is the exception to the above information. This distinctive species is not protandrous 62 Rhodora [Vol. 87 and is pollinated by halictine bees (Catling, Ph.D. thesis, U. Toronto, 1980). Most of our Spiranthes are pioneer species that can thrive in nutrient-poor soils where competition from other plants is minimal. The majority of the Ladies’-tresses are able to colonize open, dis- turbed areas such as gravelly shores and highway ditches. Some of these plants display the aggressiveness associated with “weedy” spe- cies. Apparently, there is a comparatively short interval between germination and flowering in this genus, as Ames (1921) reported flowering plants that were only three years old. This genus is taxonomically difficult in the northeast; many spe- cies are similar in morphology, and there is a wide range of morpho- logical variation within each species. Ames (1921) attributed some of this intraspecific variation to different stages of development, as one plant may have a slender few-flowered inflorescence during its first flowering season and an elongated many-flowered inflorescence the next year. Hybrids are often formed between species that share similar habi- tats and overlap in blooming dates. This phenomenon makes identi- fication difficult and should be borne in mind whenever plants appear to be intermediate between two species. There are also indi- vidual races within species that differ in chromosome counts and reproductive strategies. All of these factors contribute to a situation that led Ames (1921) to exclaim: “Spiranthes is the most perplexing orchid genus in our flora.” Key to the Species of Spiranthes in New Hampshire 1. Flowers appearing in a single-ranked spike; plants slender. . .(2) 2. Leaves linear to oblong-elliptic, sessile; lip cream-colored or WOUOW ao biatewayaraes qeeun ese denies oe Sse eek eens (3) 3. Lip cream colored to yellowish; plants of upland habitats ee ee eee re eer ee re ee (4) 4. Spike loosely flowered; lip cream-colored ............ SELeE ETRE SR OLE SER UA DONE RS 5. Spiranthes casei 4. Spike densely flowered; lip yellowish ................ ee Te ee ee 3a. Spiranthes cernua var. ochroleuca 3. Lip with bright yellow center; plants of riverside seeps... ee ee re ore 1. Spiranthes lucida 1985] Brackley — Orchids of New Hampshire 63 2. Leaves ovate, petioled; lip with green center.............. cinta Lea CMARR EER LAS Eee es 4. Spiranthes lacera 1. Flowers appearing in several-ranked spike; plants stout ... (5) 5. Lip less than 6 mm long with a bright yellow center; flowering We Ganly SUMING! «eco 36 5h5 odes cues 1. Spiranthes lucida 5. Lip longer than 6 mm, white to yellowish; flowering in late SUMIMEE: ‘CO AUEWIN ose eer ae oars ew Sa BA ote w Ses (6) 6. Lip pandurate, markedly constricted near middle, apex dilated; sepals and petals connivent into a hood .... rere rer es re ree ee 2. Spiranthes romanzoffiana 6. Lip oblong to ovate, slightly constricted near middle, gradually tapered towards apex; lateral sepals free, petals and dorsal sepal adherent ............... (7) 7. Flowers white, nodding; plants of mesic to wet sites... ee er ere 3. Spiranthes cernua var. cernua 7. Flowers cream-colored to yellowish, ascending; plants OF CTGl SINGS ok kita a se ens cesses onda Vinena eres 1. Spiranthes lucida (H. H. Eaton) Ames, Orch., fasc. 2: 258. 1908. This species, originally described by Rafinesque as Neottia plan- taginea was based on specimens found in Troy and other parts of New York. It has also been known as Spiranthes latifolia. The name lucida means “shining,” in reference to the broad, glossy leaves. COMMON NAME: Shining Ladies’-tresses FLOWERING DATES IN NEW HAMPSHIRE: late June through July. DESCRIPTION: Plant small, slender, glabrous below to sparsely pubescent above, usually about 20 cm tall. Roots fleshy. Leaves 2-4 of graduated size, clustered near the base, oblong-elliptic to elliptic- lanceolate, shiny. Inflorescence cylindrical with few to 20 flowers in 1-2 ranks, spiralled. Floral bracts lanceolate, longer than the ova- ries. Flowers: white, tubular. Sepals linear-oblong. Lateral sepals oblique, free. Lateral petals linear-oblanceolate, connivent with dorsal sepal forming a hood. Lip oblong, folded around column with a small basal callosity on each side and a bright yellow floor. Apex of lip rounded with undulating margins. Column white with 2 bright yellow pollinia. Capsule ellipsoid. 64 Rhodora [Vol. 87 CHROMOSOME NUMBER: = 22 (+2) (P. Catling, personal communi- cation). The Shining Ladies’-tresses is the earliest member of this genus to flower in New Hampshire. Most Spiranthes are late summer and fall bloomers but this dainty little orchid begins to open its tubular flowers in June. The Shining Ladies’-tresses is unique among our Spiranthes in that the leaves do not begin to wither at flowering time but remain green and functional for the duration of the growing season (Case, 1964a). Although S. /ucida is the smallest Ladies’- tresses in the state, it is the most striking in appearance. Snow-white flowers are accented by a sunshine-bright, yellow stripe on the lip that is often bordered with fine green lines. The stem is surrounded by a basal cluster of two to four bright green, glossy leaves of graduated size, the smallest one being nearest the bottom of the stem. Unlike the majority of Spiranthes species in our region, S. lucida is pollinated by halictine bees (Catling, Ph.D. thesis, U. Toronto, 1980). Because this species prefers a substrate that is neutral or calcare- ous, its distribution is limited in New Hampshire. In Vermont, where sweeter soils prevail, the Shining Ladies’-tresses grows abundantly on river banks and in wet meadows. In that state, | have seen a proliferation of plants of Spiranthes lucida growing along the side of a small stream at the edge of a pasture. In New Hampshire, there are only two known stations where the Shining Ladies’-tresses have been found, both along rocky shores of the Connecticut River just downstream from dams. The habitats are rocky, seepy areas in full sunlight. Companion plants found at these sites are, for the most part, species considered unusual for the state and are generally restricted to the same two stations. These uncommon plants include False Asphodel (Tofieldia glutinosa), Grass-of-Parnassus (Parnassia glauca) Musky Monkey-flower (Mimulus moschatus) and Jesup’s Milk-vetch (Astragalus robbinsii var. jesupt). Because of the restricted occurrence of this species, Spiranthes lucida is listed in “Rare and Endangered Vascular Plant Species in New Hampshire” (Storks and Crow, 1978). The Shiny Ladies’- tresses is limited in its re ge to northeastern North America but disjunct stations from Missw ari and Kansas have been reported 1985] Brackley — Orchids of New Hampshire 65 (Correll, 1950). It may well be that this species is overlooked in some areas because of its diminutive proportions and its preference for rather mundane habitats. GENERAL DISTRIBUTION: New Brunswick west through Ontario, south to Virginia, west to Indiana, disjunct to Missouri and Kansas. 2. Spiranthes romanzoffiana Cham. Linnaea 3: 32. 1828. Chamiso first described this species from plants collected in the lowermost valleys of Unalaska, an island in the Aleutians. He gave the orchid the name of his patron, Nicholas Romanzoff, a Russian minister of state. COMMON NAMES: Hooded Ladies™-tresses, Romanzoff’s Ladies’- tresses FLOWERING DATES IN NEW HAMPSHIRE: July through August. DESCRIPTION: Plant with long fleshy, almost tuberous roots; erect; glabrous below, pubescent above, averaging approximately 25 cm tall. Leaves 3-6, clustered at base. linear to oblanceolate, becoming reduced to bracts on the upper stem. Inflorescence a densely flow- ered cylindrical spike of 3-ranked flowers. Floral bracts thin, ovate- lanceolate, acuminate. Flowers creamy-white to yellowish-green, gaping. Sepals and lateral petals connivent into a hood. Sepals oblong-lanceolate. Lateral petals linear. Lip with minute basal cal- losities, pandurate, markedly constricted near middle. Apex of lip dilated, recurved. Capsule ellipsoid. CHROMOSOME NUMBER: 2” = 30 (Moore, 1973). The Hooded Ladies’-tresses has the widest range of all our Spi- ranthes. In North America it has a classic boreal distribution with southward extensions into the western mountains. Spiranthes romanzoffiana also occurs in Ireland, the Scottish Hebrides, and in southwest England (S. Nilsson, 1979). The “hood” of this species is formed by the complete convergence of the sepals and lateral petals, giving the plant a “two-lipped” appearance (Case, 1964a). The lip is strongly constricted in the middle and rounded at the base and apex resulting in the characteristic “fiddle-shape.” The lip is also deflected sharply downward from the middle of the flower “with the effect of a receding chin” (Morris and Eames, 1929). 66 Rhodora [Vol. 87 Because Spiranthes romanzoffiana superficially resembles the Nodding Ladies’-tresses, S. cernua, the two taxa are often confused. In comparing plants of the two species, the spike of the Hooded Ladies’-tresses is usually denser and more tapering toward the top. The 3-ranked greenish-white flowers of S. romanzoffiana are ascending, whereas in S. cernua they are nodding and pearly-white. On the lip of the Hooded Ladies’-tresses the callosities are minute, but in the Nodding Ladies™-tresses they are prominent. Flowering times and microhabitats also separate the two species. In his botanical notebook of 1891, H. E. Sargent (unpubl., depos- ited in NHA) wrote: “Found in Plymouth (Aug. 21) two or three dried up plants that I called S. Romanz. About Sept. Ist began to find plants of S. cernua. That problem ts solved.” As Sargent noted, the Hooded Ladies”-tresses is the earlier bloomer, usually beginning to flower in July and often “gone-by” at the beginning of September when the Nodding Ladies’-tresses bloom. Occasionally the bloom- ing times of the two taxa do overlap. Spiranthes romanzoffiana prefers a wetter habitat than S. cernua and commonly is found at the edges of ponds and swales. In these wet sandy or sparsely grassy areas, the Hooded Ladies’-tresses grow along with the Bog Clubmoss (Lycopodium inundatum), the Little Green Wood Orchid (Platanthera clavellata) and the tiny, flycatch- ing sundews (Drosera spp.). Often S. romanzoffiana shares a com- munity with S. cernua, the latter occupying areas that could be termed “moist” rather than “wet.” In 1940, Steiger discovered such a locality in the town of Warner in Merrimack County, New Hamp- shire. He collected several plants that appeared to be intermediate between the two species. These putative hybrids were named Spi- ranthes Xsteigeri by D. S. Correll (1941) in honor of their discov- erer. The largest hybrid of the population was a monstrosity with abnormally developed callosities and two extra abortive lips. Case (1964a) reported a large colony of the hybrid growing in Michigan's Upper Peninsula. Hybrids between S. romanzoffiana and S. lacera var. /acera were recently reported by Simpson and Catling (1978) as occurring in Ontario and Maine. Ladies’-tresses has a delightful fragrance but descriptions of the scent are varied. Morris and Eames (1929) had it almond-scented: Gibson (1905) claimed it smelled like violets; Wherry (Correll, 1950) likened it to cumarin; Summerhayes (1951) reported it to be remini- 1985] Brackley — Orchids of New Hampshire 67 scent of either Hawthorn or vanilla; while to Bingham (1939) the odor was similar to lilacs. The pollinators of this species are listed by van der Pijl and Dod- son (1966) as being bees of the genera Halictus and Chlorhalictus (Halictidae) as observed by Godfrey in the British Isles. In western North America a separate variety of Spiranthes romanzoffiana has been described as var. porrifolia (“leek-green”). This variety is distinguished by its prominent callosities and a lip that is not significantly dilated at the apex. GENERAL DISTRIBUTION: Boreal North America; Newfoundland west to Alaska, south to New York in the east, through the Great Lakes area; extending south in the western mountains to California and New Mexico. Also Ireland, Scottish Hebrides, and southwest England. 3. Spiranthes cernua (L.) L. C. M. Rich., De Orchid. Europe. Annot. 37. 1817. Linnaeus (1753) named this species Ophrvs cernua from plants collected in Virginia and Canada. The specific epithet cernua is Latin for “nodding,” referring to the appearance of the flowers. COMMON NAME: Nodding Ladies’-tresses FLOWERING DATES IN NEW HAMPSHIRE: September to early Oc- tober DESCRIPTION: Var. cernua (3). Plants quite variable, from slender, fleshy roots, erect, glabrous below, downy pubescent above, up to 45 cm tall. Leaves 3-6, glabrous, clustered at base, linear-lanceolate, acuminate, becoming bracteate above. Inflorescence a densely flow- ered spike of nodding white flowers in 2-4 ranks. Floral bracts lanceolate, acuminate. Dorsal sepal lanceolate. Lateral sepals lan- ceolate, oblique, free. Lateral petals linear-lanceolate, connivent with the dorsal sepal. Lip with prominent basal callosities, ovate- oblong, somewhat constricted at the middle, directed downward with undulating margins at apex. Capsule ellipsoid. Var. ochroleuca (3a). This variety differs from the typical in the following attributes: straw-yellow, non-dilated lip; longer, nar- rower, more tapering floral bracts; longer, more recurved callosities: ascending flowers; greater ascension and separation of lateral and 68 Rhodora [Vol. 87 dorsal sepals; leaves that are more apt to be sessile; slightly later flowering time; drier habitats (Sheviak and Catling, 1980). CHROMOSOME NUMBERS: | Var. cernua: 2n = 60; var. ochroleuca: 2n = 30: intermediate or undetermined plants: 2n = 45 (Sheviak and Catling, 1980). Probably our most common orchid, Spiranthes cernua is wide- spread throughout the state. The lucky combination of abundance and beauty makes the Nodding Ladies’-tresses one of the most pop- ular wildflowers of our region. The snow-white, fragrant flowers have a crisp crystalline appearance and are usually tightly spiralled in two to four ranks. The “nodding” impression of the blossom, which is really more or less horizontal, derives from downward deflection of the lip. The flowering spike varies in size from 10 to 45 cm and is surrounded by grass-like basal leaves. Spiranthes cernua is the last orchid to bloom during the flowering season in New Hampshire. All summer long the fields and roadsides are dominated by naturalized plants that are European in origin, such as Daisies (Chrysanthemum leucanthemum), Buttercups (Ra- nunculus spp.), and Dandelions (Taraxacum officinale). With the coming of Autumn, our native herbs display their bouquets. The Nodding Ladies’-tresses is not found in the company of tall and leafy Asters and Goldenrods, but instead grows in roadside ditches, old sand pits, and mowed, moist meadows. The most common habitat of this species is in roadside ditches where summer mowing keeps competition with grasses to a min- imum. Apparently, modern highway systems have been instrumen- tal in making this orchid widespread by providing long stretches of appropriate habitats. A roadside area where Milkwort (Polvgala sanguinea) and Gerardia (Agalinis spp.) are blooming generally means that plants of the Nodding Ladies’-tresses are likely to be nearby. At especially favored sites one can find Bottle and/or Fringed Gentians (Gentiana spp.) in the vicinity of populations of Spiranthes cernua. It takes a heavy killing frost to nip plants of the Nodding Ladies*- tresses. | have seen flowers of this orchid in Pinkham’s Grant, Coos County, completely encased in ice in the early morning. By noon, the warm late-September sun of Indian Summer had freed the plants of the ice and the flowers looked fresh and unwilted. 1985] Brackley — Orchids of New Hampshire 69 The annual appearance of this species seems to be dependent upon rainfall. During the early part of September in 1980, | was unable to find the Nodding Ladies’-tresses in its usual haunts. In mid-September the rains finally came, and the next weekend the roadsides were dotted with the white spikes. So abundant and noticeable are these plants that it is quite possible to botanize when driving along and stopping when populations of the Nodding Ladies’-tresses appear by the side of the road. Polyembryony, the production of more than one embryo per seed, was reported long ago in Spiranthes cernua by Leavitt (1901). Swamy (1948) also investigated embryo formation in the Nodding Ladies’-tresses. He found that three separate races, sexual, asexual, and intermediate, occur in this species. The predominant race (4068 out of 5000 specimens studied) is the asexual one in which 2-6 embryos per seed arise without fertilization (agamospermy). Out of the 5000 plants that he studied, 520 were a normal sexual race in which a single embryo is produced (monoembryony) as a result of sexual fusion. Swamy found that the sexual race was mostly con- fined to the New England area. The remaining 412 plants that he studied were termed “intermediates,” as in these individual plants some embryos were produced as the result of sexual fertilization while others arose by agamospermy. Further studies have shown that the polyembryonic, asexual race is a polyploid (2n = 60) and that the sexual, monoembryonic race is diploid (2n = 30) (Sheviak and Catling, 1980). 3a. Spiranthes cernua var. ochroleuca In 1899, Andrews reported a variant of Spiranthes cernua in which the flowers were a yellowish color, rather than the usual pure-white. This color form, along with other subtle morphological differences led Rydberg (Britton, 1901) to describe a new species of Ladies’-tresses which he called Gvrostachys ochroleuca. Rydberg’s species corresponds to the sexual, monoembryonic race of S. cer- nua. Ames (1906) reduced this taxon to S. cernua var. ochroleuca:’ because the one reliable character of differentiation between the two taxa Is the type of embryo present, an impractical taxonomic character. Recently Leur (1975) reinstated S. ochroleuca to species status. Sheviak and Catling (1980) attempted to illustrate through a 70 Rhodora [Vol. 87 statistical analysis of floral morphology that morphological charac- ters alone could serve to distinguish the two taxa. I] have collected hundreds of specimens of the Spiranthes cernua complex and have found very few that neatly fall into the ochro- leuca category. Many characteristics are apparently dependent upon habitat (i.e., S. cernua sometimes has ascending flowers). Two plants of the same species, one growing in the bottom of a wet roadside in full sunlight, the other growing at the top of the road- bank in partial shade, might be expected to look somewhat differ- ent. Although the two races may sometimes be distinguished by morphology, the polyploid race is usually best differentiated from the diploid race by an examination of embryos. As Correll (1950) stated, “Such ultratechnical characters are not readily usuable.” Both varieties of the Nodding Ladies’-tresses are visited by bees of the genus Bombus (Apidae) (Leur, 1975; Ames, 1921). GENERAL DISTRIBUTION: Nova Scotia west through Ontario to Minnesota, south to Florida, west to Texas. 4. Spiranthes lacera (Raf.) Raf., Herb. Raf., 44. 1833. (Slender Ladies’-tresses) A somewhat variable taxon probably best treated as a single species composed of two varieties, var. /acera and var. gracilis. Rafinesque first described this species as Neottia lacera from plants found in New York State. Six years later it was also described as Neottia gracilis by Bigelow in his Florula Bostoniensis (1824), a compendium of plants found in the vicinity of Boston. In this work Bigelow included a description of a plant he called N. gracilis var. secunda. In this plant, which was collected in Conway, Carroll County, New Hampshire, the spike was hardly twisted and the flowers more slender than the typical plant. For many years, both varieties of this species were referred to Spiranthes gracilis. In 1946, Fernald accorded specific rank to the two entities, calling the more northern plants S. /acera, and those plants more southern in distribution S. gracilis. Other botanists (Correll, 1950; Case, 1964a) felt that this complex was best treated as a single species, S. gracilis. It has been pointed out by Fernald (1946a) and Voss (1966) that Rafinesque’s name S. /acera has prior- ity over S. gracilis, and those who prefer to treat the taxon as a single species must use the earlier name. 1985] Brackley — Orchids of New Hampshire fe The name /acera means “torn,” describing the margins at the apex of the lip. The Latin word for “slender” is gracilis, referring to the very slender flowering spike. COMMON NAMES: Slender Ladies’-tresses, Green Pearl-twist FLOWERING DATES IN NEW HAMPSHIRE: July through August (occasionally earlier or later). DESCRIPTION: (var. /acera) Plants from fleshy, fasciculate roots, very slender, moderately pubescent, up to 50 cm tall. Leaves basal, 2-5, short-petioled, ovate, present at flowering time. Inflorescence an elongated, loosely spiralled or secund spike. Flowers white, dis- tantly spaced. Floral bracts ovate, acuminate. Sepals lanceolate, acute. Lateral petals linear, adherent to dorsal sepal. Lip oblong with a green stripe and small callosities. Apex of lip with finely lacerate margins. Capsule ellipsoid. Variety gracilis differs from the above in the following attributes: Plant essentially glabrous; leaves rarely present at flowering time; inflorescence strongly spiralled; flowers closely spaced. CHROMOSOME NUMBER: 2n = 30 (Tanaka and Kamoto, 1975). Disregarding the subtle varietal differences, the Slender Ladies’- tresses is a distinctive plant in our area and is unlikely to be mis- taken for another species of Spiranthes. The common name is most appropriate as this plant is so slender and elongate that it resembles a wand of grass. Spiranthes lacera is a plant that is apt to be overlooked as it often blends in with its surroundings. I have walked by a group of Slender Ladies’-tresses, oblivious to their existence, only to be astounded by their “sudden” appearance when returning along the same path. When the sunlight strikes a patch of Spiranthes lacera, the wispy little flowers look like tiny flags blowing in the wind. The blossoms have a sweet fragrance and a lip that sports a green stripe. The shape of the basal leaves resembles that of the Rattlesnake Plan- tains, but they usually begin to wither around the time of flowering. In New Hampshire, Spiranthes lacera is most often found in dry, open pastures, especially on slopes, or in open, rocky woods. | have found this orchid at the edges of ski-slopes in the mountains. The occurrence of this species is widespread throughout the state. Because of its inconspicuous nature it is probably more prevalent than collected specimens indicate. 72 Rhodora [Vol. 87 The two varieties of Spiranthes lacera are somewhat separated geographically, with var. /acera a more northern element and var. gracilis found in more southern locations. Fernald (1946a) claimed that all specimens that he examined from northern and central New Hampshire were referable to S. [var.] /acera and that S. [var.] graci- lis was only present in the southern counties of Hillsborough and Strafford. While this separation may be the general case the situa- tion unfortunately is not quite so clear-cut, as each variety occa- sionally occurs in the “wrong” end of the state. The Slender Ladies’-tresses usually blossom in July and August but a few plants have been recorded in flower during June and September in New Hampshire. On this long flowering season Bald- win (1894) commented: “Nature must be fond of the Slender Spir- anthes or she would not permit it to flourish in comparatively dry soil and to enjoy a four months lease of life.” As tiny as the flowers are, they attract many insect visitors. Robertson (1928) observed a male Long-tongued Bee, Bombus americanorum (Apidae) and a female Short-tongued Bee Calliopsis andreniformis (Andrenidae) visiting the orchid. The Slender Ladies’-tresses propagates readily from seeds, becoming abundant once it is established in an area (Case, 1964a). This species can apparently achieve the flowering state in only three to five years (Correll, 1950). Spiranthes lacera var. gracilis has been reported to hybridize with S. vernalis, in those areas where both taxa grow and flower at the same time, to produce S. Xintermedia (Catling, 1978). This hybrid is unlikely to be encountered in this state because one of the puta- tive parents, S. vernalis apparently does not grow in New Hamp- shire. As previously mentioned, S. /acera var. lacera has been reported to hybridize with s. romanzoffiana (Simpson and Catling, 1978). GENERAL DISTRIBUTION: Nova Scotia west through Ontario to Saskatchewan, south to Florida, west to Texas. 5. Spiranthes casei Catling and Cruise, Rhodora 76: 526. 1974. This is the most recently described species among the orchids of New Hampshire. Spiranthes casei was described by Catling and Cruise to eliminate the problem of a group of northern plants which were previously referred to S. vernalis. These plants are morpho- logically quite distinct from the S. vernalis that occurs in the south- 1985] Brackley — Orchids of New Hampshire 73 western United States. The species was named for the orchid specialist Frederick W. Case, Jr., author of Orchids of the Western Great Lakes Region (1964a). COMMON NAME: Case’s Orchid FLOWERING DATES IN NEW HAMPSHIRE: August to early September. DescRIPTION: (from Catling and Cruise, 1974; and Catling, 1978) Plants erect, glabrous below, pubescent above, 27-33 cm tall at flowering time. Leaves glabrous, often glaucous, oblanceolate or linear-lanceolate, reduced to sheaths below the inflorescence, ascending up to 1/3 the length of the stem, present at flowering and withering soon after. Inflorescence a spike of loosely arranged flowers, often in a single spiral, the rachis with septate, reddish- glandular hairs. Floral bracts ovate to ovate-lanceolate, acuminate, pubescent at the base. Ovary obliquely swollen on the upper side at anthesis, pubescent. Sepals lanceolate, barely upcurved. Lip obo- vate, apically truncated and occasionally notched. Base of lip cor- date. Basal callosities stout, incurved. Capsule elliptical. CHROMOSOME NUMBER: 2n = 60-70, 65-75 (P. Catling, personal communication). In one section of his discussion of New England orchids, Ames (1921) noted that the northern Spiranthes vernalis may not be com- parable to the southwestern form of S. vernalis, as originally de- scribed by Englemann and Gray. Ames postulated that the northern vernalis may be a hybrid between S. cernua and S. gracilis, as the plants in question seemed to him to be intermediate between the two. Catling and Cruise (1974) do not deny the possibility of a hybrid origin of this species but point out that the taxon maintains itself in the abscence of one or both putative parents, shows no evidence of back-cross-swamping of its distinctiveness, and is fer- tile. Additionally, Spiranthes casei has cream-colored flowers, whereas S. cernua and S. lacera both have white flowers. While plants clearly referable to Spiranthes vernalis occur in southeastern Massachusetts, these populations seem to constitute the northern limit of this species. It appears that S. vernalis (sensu Englemann and Gray) has not been found in New Hampshire to date. The specimens identified as S. vernalis from this state that | have seen are referable to S. casei, or are hybrids of uncertain 74 Rhodora [Vol. 87 affinities. Catling has placed the hybrid S. cernua X S. gracilis Ames and S. neglecta Ames in synonymy with S. vernalis. He has also indicated that the name S. Xintermedia Ames (pro hybr.) should be retained for hybrids between S. vernalis and S. lacera var. gracilis. All of the above taxa are similar to each other but differ from S. case! in the light-colored and non-glandular pubescence of the rachis (Catling, 1978). The pubescence of S. casei is a dark-reddish color, septate, and glandular. Spiranthes intermedia, as recently circumscribed by Leur (1975), is actually synonymous with S. casei. Spiranthes casei is a tall, robust plant that grows in dry open areas such as roadside banks and along power lines. I have found this species growing in an old field in Colebrook, Coos County, in August. Apparently, this orchid flowers earlier than the Nodding Ladies’-tresses with which it sometimes grows (Catling and Cruise, 1974). An examination of herbarium specimens of Spiranthes from the state turned up some plants from northern New Hampshire that easily fall into the S. casei category. The situation becomes more complicated in the southern counties as a few plants labeled “S, vernalis” appear to be neither S. vernalis nor S. casei, and are most probably hybrids. In their study of their new species, Catling and Cruise found S. casei to be partially apomictic. Flowers from which insects were excluded demonstrated 10-25% seed germination. GENERAL DISTRIBUTION: A broad band from Nova Scotia west through northern New England to Michigan and Ontario (Catling and Cruise, 1974). 10. Isotria Raf., Med. Repos. II 5: 357. 1808. This genus consists of two eastern North American species which were originally referred to Arethusa. In 1808 Rafinesque segregated these species into his genus /sorria but his generic concept did not gain acceptance until about 70 years ago. In the interim most bota- nists followed Nuttall’s (1818) treatment which included the two species within the genus Pogonia. The apparently* whorled arrangement of the leaves at the sum- mit of the stem is unique to this genus of the orchid family. The *Homoya (Master's thesis, So. Ill. Univ., 1977) reported that “careful inspection reveals staggered leaf insertion on the stem.” 1985] Brackley — Orchids of New Hampshire 75 name /sorria is from two Greek words meaning “equal” and “three,” referring to the nearly identical size and shape of the sepals. Key to the species of /sorria 1. Flowers ona pedicel over 2 cm long; sepals purplish, more than 3 cm long; stem purplish; in fruit the pedicel is about twice the lene ol he capsule oasvsnseee5e345% 1. /sotria verticillata 1. Flowers on a very short pedicel, appearing almost sessile; sepals light green, less than 3 cm long; stem greenish-white, in fruit the pedicel is about the same length as the capsule ........ (PREG Sete Ene eeedak eae eee een ces 2. Isotria medeoloides 1. Jsotria verticillata (Muhl. ex Willd.) Raf., Med. Repos. II 5: 357. 1808. Muhlenberg first collected this species in the Blue Ridge Moun- tains of Pennsylvania; he sent the specimens along to Willdenow in Germany who then published the description of the plant (as Are- thusa verticillata) in his Species Plantarum (1805). The epithet verti- cillata is Latin for “whorled,” describing the apparent arrangement of the leaves. COMMON NAME: Large Whorled Pogonia. FLOWERING DATES IN NEW HAMPSHIRE: mid-May to mid-June. DESCRIPTION: Plant erect with extremely long, hairy roots that produce root-shoots which give rise to new plants. Stem glabrous, hollow, purplish, 10-30 cm tall. Leaves 5-6, green, oblong- lanceolate, whorled at summit of stem. Inflorescence consists of | or 2 pedicellate flowers terminating the stem. Sepals purplish, narrowly lanceolate, acuminate, spreading. Lateral petals yellow- green, elliptic-obovate. Lip 3-lobed, oblong-cuneate, yellowish- white, streaked with purple, crested along median ridge. Column approximately | cm long with a denticulate apex. Capsule erect, ellipsoid, on a pedicel nearly twice the length of the capsule. CHROMOSOME NUMBER: 2n = 18 (Baldwin and Speese, 1957). The Larger Whorled Pogonia presents an exotic-looking flower. Elevated on a pedicel above the circle of leaves, the dramatically long and spreading purple sepals serve to focus attention on the yellowish-green petals which they surround. 76 Rhodora [Vol. 87 Isotria verticillata is primarily a more southern species that reaches its northern limit in New England. In the more optimum southern environments the Large Whorled Pogonia is considered uncommon but it is truly rare here at the northeastern edge of its range. The historic occurrence of this species in the southern part of New Hampshire is verified by three specimens, all collected before 1909. In the literature /. vertici/lata has been mentioned as occur- ring in Auburn and Manchester in 1900 (Batchelder, 1900). Unfor- tunately the station in Manchester at which the Large Whorled Pogonia was described by Batchelder (1900) as being locally abundant, was transformed into a potato field “doubtless more profitable to the owner of the land than the orchids, but less inter- esting to the botanist” (Batchelder, 1909). Presently there is one known extant station for this species in New Hampshire. The report of the orchid’s occurrence in Rockingham County by Storks and Crow (1978) proved to be the Small Whorled Pogonia, /sotria medeoloides. Although the Large Whorled Pogonia is found in a variety of habitats throughout its range, in New Hampshire it has been con- sistently reported as growing in second (or third) growth acidic woodland habitats dominated by Red Maple (Acer rubrum) and Hemlock (Tsuga canadensis). Companion herbs include Canada Mayflower (Maianthemum canadense), Goldthread (Coptis groen- landica) and Wintergreen (Gaultheria procumbens) “for which the Pogonia seemed to have a fondness” (Morris and Eames, 1929). Isotria verticillata very closely resembles the Indian Cucumber-root (Medeola virginiana) which is also commonly associated with the orchid. In a recent study of the breeding systems and pollination biology of the genus, Mehrhoff (1980) found that /. verricil/ata is pollinated by bees of the families Andrenidae, Anthophoridae and Halictidae. In the same study it was noted that percentage seed set in this species was only 15% owing to low pollinator activity. The Large Whorled Pogonia regularly propagates by means of root-shoots and often forms large colonies (Ames, 1922a; Leur, 1975). Isotria verticillata is reported as being subject to long periods of dormancy but Case (1964) has observed a colony of this species that has produced plants annually for 13 years. Flowering is reported to be sporadic in the Large Whorled Pogonia but this behavior can 1985] Brackley — Orchids of New Hampshire 77 probably be attributed to factors such as lack of maturity and non- optimum environmental parameters. GENERAL DISTRIBUTION: New England west to Michigan, south to northern Florida, west to eastern Texas. 2. Isotria medeoloides (Pursh) Raf. Fl. Tellur. 4: 47. 1838. This species was first described by Pursh in 1814 as Arethusa medeoloides. The Small Whorled Pogonia has also been known as Pogonia affinis and Isotria affinis. The specific epithet medeoloides means “resembling Medeola,” the Indian Cucumber-root, a com- mon liliaceous plant of the eastern deciduous forest. COMMON NAMES: Small Whorled Pogonia, Little Five Leaves. FLOWERING DATES IN NEW HAMPSHIRE: June DESCRIPTION: Plant erect, glabrous, up to 27 cm tall, elongating in fruit up to 36 cm. Roots fibrous, hairy. Stem stout, pale green, glaucous, hollow. Leaves terminal, 5 or 6 ina whorl, elliptic, apicu- late, slightly drooping when flowering begins and becoming hori- zontal in fruit. Inflorescence of 1 or 2 nearly sessile flowers, the short pedicel only becoming noticeable in fruit. Flowers yellowish- green. Sepals linear-oblanceolate. Petals oblanceolate. Lip whitish- yellow with greenish veins, three lobed. Lateral lobes triangular. Mid-lobe rounded, cuneate, with a median crest. Plant greatly elongating in fruit. Capsule erect, ellipsoid, on a short pedicel about equaling the capsule in length. CHROMOSOME NUMBER: 27 = 18 (Baldwin and Speese, 1957). “Our rarest orchid,” state Gleason and Cronquist (1975) of /sotria medeoloides in their Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Leur (1975) concurred that the Small Whorled Pogonia is the rarest orchid east of the Mississippi, excluding Florida. Many botanists have never seen this species in the wild. | am fortunate to be among those that Ames (1922) called the “favored few” who have actually found /. medeoloides. In other parts of the country reports of new /. medeoliodes stations have often turned out to be /. vertici/lata, whereas in New Hampshire the opposite situation has occurred. 78 Rhodora [Vol. 87 A review of the characters that serve to distinguish the two taxa seems appropriate here. Their very names suggest a size differential but in newly discovered populations of /sotria medeoloides in Maine and New Hampshire, many individuals exceed the height heretofore listed in published descriptions (E. Briggs, personal communication). A better indicator is the length and shape of the sepals which, in /. verticillata, are over 3 cm long, purplish in color, and are widely spreading. In /. medeoloides the sepals smaller, not spreading, and greenish-colored. The flowers of the former are borne on a pedicel but in the latter they appear to be almost sessile. In the Large Whorled Pogonia the stem is purplish while in the Small Whorled Pogonia it is a pale greenish color. When the plants are in fruit the pedicel of /. verticillata is twice the length of the capsule whereas in /. medeoloides they are about equal in length (Case, 1971). Another interesting difference between the two species is the posi- tion of the whorled leaves when blossoming is taking place. As flowering begins the leaves of the Large Whorled Pogonia are pointed upward and gradually drop to a horizontal position. In the Small Whorled Pogonia the leaves are pointed toward the ground when the flowers first open and slowly rise to become horizontal. Plants of /sotria verticillata are described as growing in clumps within a few inches of each other, while /. medeoloides is said to commonly occur as solitary plants growing several feet apart (Case, 1971). Although this report may be generally true, | have observed clumps of three to four flowering plants of the Small Whorled Pogonia growing up from almost the same point in the ground. In the past, most known populations of the Small Whorled Pogonia have consisted of very few individuals. Mehrhoff (1980) reported that in all of the populations of this species known as of 1979, the mean average number of plants per population was five, and the median average was listed as being only three. At the other extreme, some newly-located stations in N.H. and Maine have well over 200 plants. Because /. medeoloides is so rare throughout its range, it has been listed by the U.S. Fish and Wildlife Service (Fed- eral Register, Vol. 47, No. 176, Sept. 10, 1982) as “Endangered” under the Endangered Species Act of 1973. Until recently, it was believed that there were a greater number of dried specimens of this taxon in herbaria than there are living plants (Federal Register, 1985] Brackley — Orchids of New Hampshire 79 Vol. 45, No. 242, December 15, 1980). Of all the individual plants known to exist in the world today, approximately 80% occur in Maine and New Hampshire (R. Dyer, personal communication). Isotria medeoloides grows in acidic soil conditions and is most often found in second growth mixed deciduous forest communities. The preferred habitat appears to be in areas of thick organic duff associated with vernal streams. In other words, this species may be found in a forest habitat typical of many areas of New Hampshire. Two small sub-populations of /sotria medeoloides have recently been found in dark, fairly wet, coniferous forests in New Hamp- shire, but none of the plants flowered in that situation. It has been claimed that the Small Whorled Pogonia remains dormant underground without flowering for up to 20 years, but this supposition has not been supported by detailed observations. One New Hampshire station has been observed since 1965 by Mr. and Mrs. Ring, a naturalist couple in Merrimack County. They have made a valuable contribution to the knowledge of the biology of this species by keeping careful records of its occurrence. Though the number of flowering plants varied from year to year, the population made a showing every year. When the plants were discovered, the area (which had been previously logged) was composed mainly of saplings about 12 feet tall and a few scattered larger trees. The Rings have found that the percentage of plants blossoming increased between 1965 and 1973. Since that time the number of flowering individuals has steadily declined. These data seem to sug- gest that light intensity may provide at least one stimulus for flower- ing. These observations have been reinforced by Mehrhoff’s recent work on the genus. He found that flowering individuals of the Small Whorled Pogonia occurred in locations with only a low herba- ceous cover while sites with relatively dense shrub coverage, or many saplings, only produced vegetative individuals (Mehrhoff, 1980). One station of Small Whorled Pogonia that I located in 1980 consisted of only two small plants. In 1981 this area was hard hit by an invasion of Gypsy Moths which had devoured much of the canopy. The population has doubled in size and vigor each year since then, a phenomenon that I believe to be related to the increased amount of sunlight reaching the forest floor. Lacking scent, nectar guides and brightly colored flowers, the Small Whorled Pogonia has been found to be self-pollinating by 80 Rhodora [Vol. 87 Mehrhoff (1980) who also noted that seed set was 83% in this species. Despite this relatively high percentage of seed set, a study of an Illinois population suggests that /. medeoloides exhibits reduced reproductive potential owing to early termination of capsule devel- opment and late summer mortality, at least at that particular site (Homoya, Master’s thesis, So. Ill. Univ., 1977). | have also observed many cases of arrested flower development in this species, where flower buds were formed but eventually withered away. That this orchid is exceedingly rare is indisputable, but its pres- ence also just may be overlooked much of the time. A small green plant that resembles many other herbs and lives in an ordinary, unspecialized habitat (where few botanists concentrate their field work) can easily go unnoticed. Perhaps the secret of its continued existence, despite the low numbers of known populations which would seem to indicate imminent extinction, lies in the fact that the Small Whorled Pogonia is able to avoid detection by would-be exploiters. DISTRIBUTION: Ontario, Canada; Connecticut, Georgia, Illinois, Massachusetts, Michigan, Missouri, New Hampshire, New Jersey, New York, Maine, Maryland, North Carolina, Pennsylvania, Rhode Island, South Carolina, Vermont and Virginia. 11. Pogonia Juss. Gen. Pl. 65. 1789. Pogonia is a small genus of acid-loving orchids which occurs in eastern North America and eastern Asia. The first-described species of this genus was originally included within the genus Arethusa by Linnaeus (1753); de Jussieu transferred a section of Linnaeus’s genus Arethusa to his new genus Pogonia. Several species were formerly referred to Pogonia but are now segregated in distinct genera, including Cleisres, Isotria, Triphora and Psilochilus (Ames, 1922a). In this genus the pollen grains of the pollinia characteristically are separate (simple) rather than being grouped into tetrads. The name of the genus is taken from the Greek word for “beard,” referring to the bristles on the lip. 1. Pogonia ophioglossoides (L.) Juss. Gen. Pl. 65. 1789. The only eastern North American species in this genus is the Rose Pogonia, Pogonia ophioglossoides. The specific name ophioglos- 1985] Brackley — Orchids of New Hampshire 8] soides means “like the Adder’s-tongue,” referring to the plant’s sin- gle leaf which is reminiscent of the solitary leaf of Ophioglossum, the Adder’s-tongue fern. COMMON NAMES: Rose Pogonia, Snake-mouth, Adder’s-tongue Orchid FLOWERING DATES IN NEW HAMPSHIRE: Mid-June through July DESCRIPTION: Plant slender, glabrous, 8-30 cm tall. Stem green, arising from fibrous roots. Leaf solitary, fleshy, ovate-lanceolate, almost half-way up stem. Floral bracts leaflike, oblong-lanceolate. Ovary sessile, slender. Flowers terminal, usually solitary (occasion- ally 2), rose to pink (rarely white, in forma a/biflora), fragrant. Sepals narrow, linear-oblong to elliptic. Lateral petals wider than the sepals, oblong-elliptic, obtuse. Lip spatulate, narrowing at base, fringed along margins at apex, bearded along the three main veins with fleshy yellow bristles which become longer and fuchsia-colored at the apex. Column approximately | cm long, pink, curved, toothed at apex. Capsule ellipsoid, ribbed, erect. CHROMOSOME NUMBER: 2n = 18 (Moore, 1973). Our eastern North American bog-orchid, Pogonia ophioglos- soides, was brought to the attention of the world when Ker pub- lished a description and drawing of the species in The Botanical Register “consisting of Colored Figures of Exotic Plants cultivated in British Gardens with their history and Mode of Treatment” (1916). The slender plant, bearing a beautiful pink blossom with an elaborately fringed lip, understandably captured the fancy of Eng- lish nurserymen. The acid-loving Rose Pogonia occurs quite frequently in our region in wet meadows and open bogs. In the former habitat the delicate pink flowers seem to magically appear amoung the grasses. Robert Frost (1964), in his poem “Rose Pogonias” described such a scene: “For though the grass was scattered Yet every second spear Seemed tipped with wings of color That tinged the atmosphere” The presence of P. ophioglossoides is more noticeable in sphag- num bogs where many vegetative plants, consisting only of solitary 82 Rhodora [Vol. 87 leaves on small petioles, rise straight up from the bog mat. The Rose Pogonia can reproduce vegetatively by rootshoots and is able to form large colonies in favorable habitats. The fragrance of the Rose Pogonia has elicited a wide variety of descriptions. Thoreau (1884) found the odor offensive and declared that it smelled “exactly like a snake.” Others have likened its per- fume to raspberries (Barker, 1931) or violets (Baldwin, 1894). I have found the Rose Pogonia to possess a sweet odor that smells unique- ly like Rose Pogonia. The structure of the flower of Pogonia ophioglossoides 1s adapted for outcrossing (Scudder, 1862). The pollen-masses in this species are located within a pouch-like recess, covered by a hinged lid (Bingham, 1939). To reach the nectar that is located deep within a narrow passage-way, a visiting insect must first press its head against the sticky stigma. Any pollen that the bee carried into the blossom is thus removed and, at the same time, its head is smeared with the sticky substance. In the process of backing out of the small opening the head of the insect catches the projecting edge of the lid. As the bee continues to back out, the lid raises to expose the pollen- masses, parts of which adhere to the front of its head. When the bee has completely emerged the anther lid snaps shut again (Scudder, 1862). Heinrich (1979) found that the odor, fragrance, and presence of nectar in P. ophioglossoides attracts bees of the genus Bombus (Apidae), and they serve as pollinators for the orchid. The percent- age of seed set in this species is quite variable, ranging from 10-100%. The higher figure has led to speculation that some popu- lations of the Rose Pogonia may be apomictic (Thein and Marks, 1972). GENERAL DISTRIBUTION: eastern North America; Newfoundland, west through the Great Lakes region to Minnesota, south along the Coastal Plain to Florida, west to Texas. Closely related to P. japon- ica of eastern Asia (Li, 1952). 12. Arethusa L. Sp. Pl. 950. 1753. In Greek mythology Arethusa was a beautiful river nymph. Gro- novius appropriately applied the name to this orchid in his pre- Linnaean publication Flora virginica (1743). Linnaeus subsequently used the same name, Arethusa bulbosa, in his Species Plantarum. 1985] Brackley — Orchids of New Hampshire 83 |. Arethusa bulbosa L. Sp. Pl. 950. 1753. Arethusa bulbosa is the only species within the genus. A closely related Japanese plant Eleorchis japonica (Sawa-ran) has some- times been treated as Arethusa japonica (Ohwi, 1965). The specific epithet bu/bosa refers to the bulb-like corm from which the plant arises. COMMON NAMES: Arethusa, Dragon’s-mouth FLOWERING DATES IN NEW HAMPSHIRE: late May through June DESCRIPTION: Plant slender, arising from a bulbous corm, glab- rous, scapose, 10-40 cm tall. Leaf solitary, grass-like, enclosed ina sheath at flowering time and emerging after the flower fades. Floral bracts scalelike, triangular, minute. Ovaries sessile, erect. Flowers terminal, solitary (rarely 2), deep rose (rarely white as in forma albiflora, or lavender in forma subcaerulea). Dorsal sepal oblanceo- late, connivent with the linear-oblong lateral petals, all curved over the column. Lateral sepals oblong-lanceolate, oblique, only slightly curved foreward. Lip pale pink streaked with purple, clawed, oblong, recurved, often indistinctly 3-lobed. Margins of lip crenulate-undulate, erose. Disks of lip with yellow, fleshy, fringed crests. Column pink, flattened, linear-spatulate, erose at apex. Cap- sule ovoid, erect, approximately 2 cm long. CHROMOSOME NUMBER: 2n = 44(Thein and Marks, 1972). All who have had the good fortune to see this orchid in bloom acclaim its beauty. Morris and Eames (1929) stated: “To our mind this is the most exquisitely beautiful of all single-flowered orchids.” With all the orchids in the United States and Canada to choose from, Dr. C. A. Leur selected photographs of Arethusa to grace the cover of his recent book The Native Orchids of the United States and Canada excluding Florida. This colorful denizen of bogs and wet meadows is sometimes called “Dragon-mouth,” for its animal- like countenance. The distribution of Arethusa is restricted to northeastern North America with one disjunct population in the mountains of North Carolina. In New Hampshire the number of known populations of this striking orchid has significantly decreased since the turn of the century. Fernald (1950a) stated that A. bu/bosa is “rapidly be- coming extinct S. of Nfld. and Canada.” 84 Rhodora [Vol. 87 Reasons for the drastic reductions of our Arethusa populations are circumstantial; unlike most orchids in our flora, Arethusa seldom reproduces vegetatively and depends primarily upon seed produc- tion to perpetuate itself (Case, 1964a). In addition, the bulb-like corm is only loosely attached in the mossy substrate where the slightest disturbance may dislodge it (Fernald, 1950a). In Rhode Island, Stuckey (1967) noted that the “changing patterns of land use away from pasturing and mowing would seem to be the direct cause of the decline in numbers of Arethusa. Some of the old Aretfhusa stations are no longer suitable habitats for the orchid due to the natural tendency of wetlands in our area to become drier over a long period of time (Jorgensen, 1971). One Arethusa location in Carroll County prompted Wolfboro botanist H. E. Sargent to write in his botanical notebook, in 1917, “never saw so much of it” (Sargent, unpublished notebook, NHA). During the summer of 1980 the same location yielded only two Arethusa blossoms in the now drier meadow. An aesthetically pleasing plant like Arethusa is unfortunately vulnerable to the possessiveness of man; many hundreds of these orchids have been taken from their boggy habitats only to perish in wildflower gardens or plant presses. Arethusa attracts pollinators to its showy, fragrant flowers which produce a small amount of nectar. Thein and Marks (1972) found that A. bu/bosa requires a large insect to effect pollination, and this role is usually filled by queens of the genus Bombus (Apidae). The mechanism of pollination is similar to that of Pogonia. In this case it is the bee’s thorax that comes in contact with the sticky stigma, forces the anther lid open, and receives the (usually whole) pollinia. Percentage seed-set is quite low for A. bu/bosa, approximately 5% in northern Wisconsin (Thein and Marks, 1972). Baldwin (1894) quoted “the unsentimental Hooker” (Sir William Jackson Hooker) as stating that Arethusa bulbs are used as a cure for toothache. In our era of preventive dentistry one hopes that the corms of this striking orchid are safe from such cure seekers. GENERAL DISTRIBUTION: northeastern North America; Newfound- land west through the Great Lakes region to Wisconsin, south to Pennsylvania, disjunct to North Carolina. 13. Calopogon R. Br. /n W. et W. T. Aiton. Hort. Kew ed. 2, 5:204. 1813. (conserved name). 1985] Brackley — Orchids of New Hampshire 85 Originally named Limodorum by Linnaeus (1753), this genus was removed to Ca/opogon by British botanist Robert Brown. There are four species in this genus, of which 3 are confined to the south- eastern United States. The fourth species, C. tuberosus, is more widespread. Ca/opogon is distinct in having non-resupinate flowers. In this genus the lip occupies the uppermost position in the blos- som. The name Ca/opogon means “beautiful beard” referring to the bristles on the lip. 1. Calopogon tuberosus (L.) BSP, Prel. Cat. N. Y. 52. 1888. The Grass-pink was named by Britton, Sterns and Poggenberg (abbreviated BSP). This species has also been known as Calopogon pulchellus, a most descriptive appelation meaning “little beauty.” COMMON NAMES: Grass-pink, Calopogon FLOWERING DATES IN NEW HAMPSHIRE: mid-June through July DESCRIPTION: Plant arising from a small ovoid corm with many fibrous roots. Scape slender, glabrous, 9-50 cm tall. Leaves | (rarely 2), grass-like, basal, linear, ribbed. Inflorescence a loosely- flowered terminal raceme of 1-8 successively opening blossoms. Flower buds sickle-shaped. Floral bracts small, approx. 6 mm long, ovate-lanceolate, acuminate. Flowers deeply colored magenta or rose-purple (rarely white). Middle sepal oblong-elliptic, acute. Lateral sepals ovate, oblique, acute, concave. Lateral petals short- clawed, oblong-pandurate, narrower than the sepals. Lip upper- most, linear with 2 basal side knobs (obscure lobes), dilated into a triangular or rounded “whale-tailed” apex, retuse. Disk bearded along the 3 veins with spreading clavellate hairs that are white with yellow tips. Column incurved, winged, spoon-shaped. Capsule ellipsoid, erect, ribbed. CHROMOSOME NUMBER: 2n = 40 (Thein and Marks, 1972). It is always a pleasant surprise to find a population of the deeply-colored Grass-pinks in bloom. To describe more accurately such intense color, the common name of this orchid should be “Grass-magenta” or “Grass-rose-purple.” There have been other complaints concerning the names of orchids. “Pogonia! Calo- pogon!!” grumbled Thoreau (1884) in his journal entry for July 7, 86 Rhodora [Vol. 87 1852, “They would blush still deeper if they knew what names man had given them.” Although this showy plant is abundant in many places within its range it is, unfortunately, yet another orchid that appears to be declining in New Hampshire. I have never found Calopogon grow- ing in any other habitat than a quaking spruce-tamarack bog. There, among the cranberries and carnivorous Pitcher-plants the grass-like leaves and bright flowers of Ca/opogon are found, most often growing alongside Rose Pogonias. In those rare sites where Arethusa also grows, the three orchids provide a succession of rose- colored flowers from late May through July. Calopogon tuberosus is pollinated by bees of the genus Bombus (Apidae) (Thein and Marks, 1972). When the bee alights on the lip its weight causes the hinged-like lip to fall downward, lowering the insect onto the column. In this way the bee’s back comes in contact with the sticky stigma and then the pollen-masses of the anther. When the insect flies away the lip springs back up to its original position. Bombus bees are the proper weight and proportion to effect pollination in Ca/opogon whereas smaller insects are not heavy enough to cause the lip to drop. Without scent or nectar, one might ask how the Grass-pink can attract pollinators. It is believed that the yellow bristles (beard) found on Calopogon, Arethusa, and Rose Pogonia mimic the yel- low anthers of non-orchid flowers that serve as a food reward for insects (Dressler, 1981). Indeed, like anthers, the beards of these three orchids show ultraviolet reflectance patterns that are impor- tant for orientation of bees on flowers (Thein and Marks, 1972). The co-occurrence of the three orchid species in the same habitat with staggered but overlapping blooming times (first Arethusa, then Rose Pogonia and finally Calopogon) leads bees to expect that the Grass-pink will also be a source of nectar. Heinrich (1979) has found that where plant species with similar flowers bloom in close succession of each other, the first to blossom has more nectar than those following. Dressler (1981) has termed the pollination tactic of Calopogon a case of “empty promises.” GENERAL DISTRIBUTION: southeastern Canada; Newfoundland west through Ontario. Eastern U.S. west to Minnesota and Texas; Cuba and the Bahamas. 1985] Brackley — Orchids of New Hampshire 87 14. Malaxis Soland., Prodr. Veg. Ind. Occ. 119. 1788. A widely distributed genus of about 200 species most of which are concentrated in Asia and Oceania (Correll, 1950). All the species are small and uncommon. The name Malaxis is Greek for “soften- ing,” probably referring to the texture of the leaves. Only two spe- cies occur in our area. Our plants have also been referred to the genera Microstylis (“little column”) and Achroanthes (“colorless flowers”), two appropriately descriptive names. Key to the species of Malaxis in New Hampshire 1. Leaf sheathing base of stem; flowers in a loose raceme with pedicels less than 3 mm long; mid-lobe of lip long and pointed QU ONES fuck c cc cas 1. Malaxis monophylla var. brachypoda 1. Leaf clasping stem almost half-way; flowers in a dense raceme with slender pedicels 3 mm or longer; apex of lip cleft into 2 triangular lobes with a minute tooth in the sinus .......... ind dean ee eee ee Gene nee eee tea we ee ee 2. Malaxis unifolia 1. Malaxis monophylla (L.) Sw., Kongl. Svens. Vetens. Nya Hand. 21: 234, t. 3, fig. P. 1800. This species was described by Linnaeus (1753) as Ophrys mono- phyllos from type material collected in the swampy forests of Prus- sia. In 1830 Lindley transferred the species to the genus Microstylis, where it is still retained by some workers (see Moore, 1980). Our eastern North American plant was described in 1835 as a separate species Microstylis brachypoda by Asa Gray, who subse- quently changed his concept of this taxon and viewed it as a variety. In 1926 Fernald (1926a) reinstated the taxon to species status as Malaxis brachypoda, a synonym by which it is most well known. Morris and Eames (1929) designated the taxon as Malaxis mono- phylla var. brachypoda, a name that has only recently gained wide acceptance. The eastern North American var. brachypoda has resupinate flowers, thus differing from the European var. monophylla in which the pedicels twist a full 360 degrees, returning the lip to an upper- most (“hyper-resupinate”) position (Dressler, 1981). The epithet monophylla means “single-leaved”, brachypoda signifies “short- pedicelled.” 88 Rhodora [Vol. 87 COMMON NAME: White Adder’s-mouth FLOWERING DATES IN NEW HAMPSHIRE: mid-June to mid-July. DESCRIPTION: Plant small, less than 20 cm, from an ovoid corm covered with leaf-sheaths. Leaf solitary, broad, ovate-elliptic, with a long sheathing petiole. Raceme slender, elongate, loosely flowered. Floral bracts minute. Flowers tiny, pale greenish-white to yellowish- green, on short, thread-like pedicels. Sepals linear-oblong, acumi- nate. Lateral petals linear, acute, reflexed. Lip lowermost, 3-lobed, slightly cordate. Lateral lobes small, rounded, inrolled. Mid-lobe longer, lanceolate, sharply pointed. Column short, stout. Capsule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 28 (Moore, 1973). Malaxis monophylla is a circumboreal species that probably sur- vived the Ice Age close to the margins of the glacier (Correll, 1950). Accordingly, our variety prefers cold wet soils and doesn’t venture further south than the glacial boundary. In Europe M. monophylla is extremely rare, occurring in calcareous swamps under dense growths of alders (S. Nilsson, 1979). The White Adder’s-mouth has the distinction of possessing the smallest flowers of all our native orchids. The tiny, pale flowers are almost colorless or “faded like withered tissue” (Morris and Eames, 1929). It is so well camouflaged by its surroundings that it is surpris- ing that this species is ever found by orchid hunters. The majority of specimens of Malaxis monophy'la that I have seen were collected in fruit. The light brown capsules are evidently more noticeable that the almost colorless flowers. The habitat preference of Malaxis monophylla var. brachypoda is reported to be wet, swampy woods, especially of Cedar (Thuja). Case (1964a) suggested that the simplest way to locate this orchid is to search in the very wettest deer trails and rabbit runs. This variety seeks the shade of the forest in this state, but toward the northern edge of its range the White Adder’s-mouth can be found growing in open areas. I was recently surprised to come upon this species in a decidedly acidic mixed-forest along the edge of a vernal stream. Malaxis monophylla var. brachypoda has been verified from only three places in the state. Because of its scarcity it is regarded as an endangered species in New Hampshire by Crow and Storks (1980). 1985] Brackley — Orchids of New Hampshire 89 On a regional level the White Adder’s-mouth'is classified as “Rare” in New England (Crow er al., 1981). The members of this genus appear to be adapted for insect polli- nation. Ina Malaxis species from India, Hooker found a projection which, when touched, turned into a drop of viscid material (Gibson, 1905). Hagerup (1951) reported that the exclusion of insects of another small member of this genus, M. pa/udosa, prevented the formation of seeds. Although I have found no record of specific pollinators for Malaxis monophylla, it may well be that small flies and gnats effect pollination in this species, as has been long sus- pected by other workers. On the European continent the common name of this orchid is “Gnat Flower” (S. Nilsson, 1979). GENERAL DISTRIBUTION: Newfoundland south to Pennsylvania, west through the Great Lakes region to Minnesota and Manitoba, north to Alaska. 2. Malaxis unifolia Michx., Fl. Bor. Am. 2: 157. 1803. This species was discovered by Michaux and described in his Flora Boreali-americanae (1803). The plants on which he based his description were collected in wooded areas of Pennsylvania, Caro- lina, and Florida. This plant has also been known by the specific epithet ophioglossoides, meaning “like the Adder’s-tongue fern.” The name unifolia refers to the single leaf produced by this species. COMMON NAME: Green Adder’s-mouth FLOWERING DATES IN NEW HAMPSHIRE: July through August. DESCRIPTION: Plant erect, glabrous, bright green, up to 27 cm tall in our area. Stem arising from a basal corm which is covered by leaf sheaths. Leaf solitary, sheathing stem about half-way, ovate, dark green. Inflorescence a densely flowered raceme of long pedicelled green flowers loosely arranged below and crowded near summit. Floral bracts minute, triangular. Sepals filiform, strongly reflexed, curled under lateral sepals. Lip auricled, oblong-quadrate, divided at apex into 2 triangular lobes with a small tooth in the open sinus. Column a minute stub. Capsule ellipsoid. CHROMOSOME NUMBER: unknown 90 Rhodora [Vol. 87 Malaxis unifolia has an unusual distribution. It occurs in eastern temperate North America, Jamaica, Cuba, and in sections of Cen- tral America. The Green Adder’s-mouth is yet another orchid so small and inconspicuously flowered that it led Baldwin (1894) to complain that he “scarcely knew how to describe or make my finest pointed pencil flatter.” With a magnifying glass and some imagina- tion one can perceive in the flower the flattened head of a viper from which the fanciful common name is drawn. In our range the Green Adder’s-mouth is adapted to a wide variety to habitats including rich deciduous woods, wet woods, moist pastures, and dry rocky slopes. Wherry (1918) stated that M. unifolia prefers soils that are acid in reaction, a requirement readily fulfilled in New Hampshire as demonstrated by the many specimens collected in the state. In compiling the records for this plant I was struck by the abundance of older collections; almost half of them are from the 19th century. Of the 85 New Hampshire specimens (many of them from the same town) deposited in New England herbaria, only 15 were collected in the past 50 years. It is difficult to assess the situation with regard to the dearth of modern collections of Malaxis unifolia. It is possible that the numbers of this species have declined owing to changing land use patterns in the state. In Rhode Island Stuckey (1967) recorded a stand of Malaxis unifolia growing with other orchids under some power lines. During a three-year period the amount of woody plants increased and the stand of orchids declined, leading her to conclude that the loss of orchid plants was caused by competition from the woody species. Similar changes have taken place in New Hampshire where many formerly open areas have reverted to forest lands during the past 100 years. It is also probable that the number of botanists working in the state, especially serious “amateurs,” has declined. This loss has greatly influenced the amount of botanical collecting now going on in New Hampshire. For the most part these individuals earned their livings in other professions but were able to spend long summer holidays in New Hampshire, where they actively pursued their avo- cation of botany. Without such collectors as Walter Deane, C. F. Batchelder, E. F. Williams and many others, our flora would be considerably less well represented in the major herbaria of the area. 1985] Brackley — Orchids of New Hampshire 91 My first experience with the Green-Adder’s-mouth came on a collecting trip specifically for ferns. The day was quite warm and the myriad of old logging roads kept us moving in circles instead of heading for the cliffs we hoped would yield some “good” pterido- phytes. The forest floor was carpeted with plants of Canada May- flower (Maianthemum canadense) and, sitting down to take a sip of water, I noticed one “Maianthemum” that looked a little different. Closer inspection revealed the plant to be the tiny Green Adder’s- mouth, a single orchid in a sea of Canada Mayflower. It is often characteristic of orchids to grow with “look-alike” plants, non- orchid species that have a similar morphology. There apparently has been no study of the pollination biology of Malaxis unifolia. Seeds are regularly produced in the Green Adder’s-mouth, an occurrence that seems to initiate the lengthening of the stem, for plants with seed capsules are generally much taller than flowering plants (Correll, 1950). GENERAL DISTRIBUTION: Newfoundland west to Ontario and Minnesota, south to Florida, Cuba, and Jamaica, west to the Gulf Coast of Texas; Mexico, Honduras, and Guatemala. 15. Liparis L. C. M. Rich., De Orchid. Eur. Annot. 38. 1817. (con- served name.) A large genus of approximately 250 species distributed through- out temperate and tropical regions, Liparis is represented in eastern North America by only two species. The genus is characterized by its fleshy, oily-looking leaves, a feature that gave the genus its name; Liparis derives from a Greek word for “greasy.” Linnaeus (1753) originally placed those members of this genus which he described into the genus Ophrys. Liparis is closely related to the genus Malaxis and can be distinguished from that genus by its elongated column. Liparis lilifolia, the Lily-leaved Twayblade, is a plant of the east- ern deciduous forest. Batchelder (1909) reported the presence of this species in Manchester, Hillsborough County, N. H., and this record has been documented by a specimen collected by him. Since this orchid no longer appears to be part of our flora, L. /ilifolia is not included in this treatment of the orchids of New Hampshire. 92 Rhodora [Vol. 87 1. Liparis loeselii (L.) L. C. M. Rich., Orchid. Europ. Annot. 38. 1817. This species was described by Linnaeus (1753) as Ophrys Loeselii, from plants collected in the swamps of Sweden and Prussia. Lin- naeus named the plant for Loesel who originally described this species in his flora of Prussia. COMMON NAMES: Fen Orchid, Loesel’s Orchid FLOWERING DATES IN NEW HAMPSHIRE: mid-June through July. DESCRIPTION: Plant from a bulbous corm, erect, glabrous, fleshy, up to 16cm tall. Leaves 2, basal, sheathing corm and stem, oblong- elliptic, keeled beneath. (Leaves reduced, broader in Open situa- tions.) Inflorescence a slender raceme, few flowered, with minute floral bracts. Flowers yellowish-green on thread-like pedicels. Sepals lanceolate, slightly recurved. Lateral petals linear, curved, spreading. Lip narrowly-cuneate to oblong, arcuate-recurved with an apiculate apex. Column stout, approximately 2 mm long. Cap- sule ellipsoid, ascending. CHROMOSOME NUMBER: 2n = 32. Liparis loeselii is a small, inconspicuous plant, often only attain- ing a height of 6 cm in this area, although more robust forms may grow as tall as 16 cm. The base of the stem is expanded into a bulbous corm, sometimes termed a pseudobulb, a feature common to tropical tree-dwelling orchids (Summerhayes, 1951). Usually the bulb from the previous year is still apparent at the bottom of the present year’s flowering shoot (S. Nilsson, 1979). The yellowish- green flowers are rather curious looking with their widely spread- ing, thread-like petals and sepals. When growing among grasses, as this plant often does, the Fen Orchid is difficult to spot. The shiny, somewhat greasy-looking leaves vary in form depending on the habitat. In shady, moist situations the leaves tend to be longer and narrower while in plants growing in drier, more Open areas the leaves tend to be shorter and broader. The common name “Fen Orchid” is European in origin and is descriptive of the habitat in which this species grows. A fen is a wet area that is neutral or alkaline in nature (Summerhayes, 1951). In 1985] Brackley — Orchids of New Hampshire 93 New England Liparis loeselii is often found growing in circumneu- tral grassy pastures and damp meadows, It is unusual among the orchids of our flora in that it also flourishes in sterile, sandy drain- age areas. I found this species in 1972 in a sandy seepage depression on the banks of the Connecticut River. Here, with little competition from other vegetation, a small population of flowering plants grew scattered among the rocks. When I returned to this station in 1978 only a few vegetative individuals were seen. In 1980 the plants were nowhere in evidence. Since Liparis /oeselii cannot withstand a hot, dry habitat, the population was probably a victim of the recent drought-like situation to which the state has been subjected. Storks and Crow (1978) listed Liparis loeselii as “widespread but infrequent” in New Hampshire. In this state the Fen Orchid has most often been found in Coos County or along the lower Connec- ticut River Valley, indicating its preference for neutral soils. This orchid certainly must be frequently overlooked because of its small size and overall “greenness.” The inconspicuous flowers of the Fen Orchid have no odor and produce no nectar (Kirchner, 1922). Hagerup (1941) studied the reproductive biology of Liparis loeselii in Denmark and found the species to be autogamous. His experimental plants were sheltered from insects but still developed seeds. In his description of the process of self-pollination he reported that the anther cap lifts up to expose the pollen-masses and subsequently collapses, forcing the pollen-masses down onto the stigma. More recently Catling (1980b) carried out a series of pollination experiments with plants of Liparis loeselii from Ontario. He observed no insect visitors in the field and determined through experiments that the North American plants are also self-pollinat- ing. In addition, Catling demonstrated experimentally that rain- drops greatly increase the incidence of autogamy, a fascinating adaption to the cool, rainy climates which best support this orchid. GENERAL DISTRIBUTION: Central Europe and northeastern North America, Nova Scotia west through the Great Lakes region to Ontario, disjunct to Saskatchewan and Washington, south to the mountains of North Carolina and disjunct to the Piedmont Plateau of Alabama. Closely related to L. hawaiensis of Hawaii (S. Nilsson, 1979). 94 Rhodora [Vol. 87 16. Calypso Salis. Par. Lond. Pl. 89. 1807. (conserved name). Because the single species in this genus possesses a sac-like lip, Linnaeus (1753) included it within the genus Cypripedium. It is now believed that the two genera evolved quite independently and are not closely related. Ca/ypso appears to represent a unique line of development in the family. In 1807 Salisbury placed this species in a separate genus and named it for the sea-nymph of Homer’s Odyssey. 1. Calypso bulbosa (L.) Oakes in Z. Thompson, Hist. Vermont 1: 200. 1842. Originally described from plants of Lapland, Russia and Siberia, this orchid is circumboreal in distribution. At various times this species has been called Calypso borealis and Cytherea bulbosa. Oakes made the presently accepted combination in Thompson’s History of Vermont (1842), a rather unusual avenue of botanical publication. The eastern North American representative has been treated by Leur (1975) as var. americana. The epithet bu/bosa is Latin for “bulbous,” referring to the bulb-shaped corm (sometimes termed a pseudobulb) at the base of the stem. COMMON NAMES: Calypso, Fairy Slipper FLOWERING DATES IN NEW HAMPSHIRE: mid-May to early June. DESCRIPTION: Plant erect, glabrous, from a bulbous corm, up to 16 cm tall. Stem purplish, glaucous, enveloped about half-way by 2 or 3 transparent sheaths. Leaf from top of corm, petiolate. Blade ovate to cordate, plicate, dark bluish-green. Floral bract lanceolate, pink. Flower showy, solitary, on a slender pedicel. Sepals and lat- eral petals linear-lanceolate, spreading, pinkish-purple. Lip saccate, pendant, with a bicornuate apex, purplish. Margins of sac folded outward forming a whitish apron. Apron spotted with purple, bearded with yellow bristles in the center. Column petaloid, convex, semiorbicular, inverted over opening of lip. Capsule erect, ellipsoid, approximately 2 cm long. CHROMOSOME NUMBER: 27 = 28. The exquisite beauty of this orchid is renowned. The story is often told of the great naturalist John Muir who, upon finding 1985] Brackley — Orchids of New Hampshire 95 Calypso for the first time, shed tears of joy over its beauty. Correll (1950) characterized Calypso as “the most beautiful terrestrial orchid in North America.” Morris and Eames (1929) penned the following: The extraordinary delicacy of the whole blossom, the miraculous blend of so many diversly gay colors, the structure of the lip, its multiplicity of detail and unique form, serve to create a thing of beauty unmatched in all of the world of flowers. In the northeast Calypso is most often found in dark, mossy woods of Northern White Cedar (Thuja occidentalis) and Spruce (Picea). There it often grows in the company of Creeping Snow- berry (Gaultheria hispidula) and the flower so favored by Linnaeus, the delicate, trailing Twinflower (Linnaea borealis). My only vision of the diminutive Calypso was in such a habitat in Vermont, where the orchid was found growing on a:moss-covered log. Despite hav- ing seen many herbarium specimens of Calypso, I was still unpre- pared for its diminutive proportions and bright coloration. One of the earliest of our wildflowers to bloom, Calypso is unaf- fected by early spring frosts of Northern New England (Nylander, 1935). Outside North America this species tolerates the cold cli- mates of Sweden, Finland, northern Russia, and northern Asia (S. Nilsson, 1979). Through most of its range Calypso is rare and local. Only in some areas of western North America can Calypso be termed “rather frequent” (Correll, 1950). It is in these western loca- tions that the plant attains a larger, more robust form which has been referred to as var. occidentalis Leur (Leur, 1975). In the three northern New England states, which delineate the southeastern range limit of Calypso, the orchid is considered “Endangered / Threatened” (Crow er a/., 1981). Of the three, the state of Maine boasts the most stations for the showy little plant. In the past Calypso was locally abundant in many localities in Ver- mont, but at the present time populations of this orchid have dwindled to just a few extant sites in the northeastern section of the state (Countryman, 1978). In New Hampshire Calypso has been found at only two locations. There is a record from Hanover, Grafton County, dated 1893 but it is unlikely that this population still exists. Our only other record is from Columbia, Coos County. In 1931, Thomas W. Wallace, a 96 Rhodora [Vol. 87 native of Columbia, discovered Calypso at the edge of Lime Pond in that town (Wallace, unpubl. letter in NHA files). He sent a photograph of the plant to Arthur Stanley Pease, author of the Flora of Coos County, who at that time was president of Amherst College in Massachusetts. Pease went to the site in 1946 and made a collection of Calypso, but when he returned to the same spot in 1952, and again in 1961, the orchid was not in evidence (Pease, 1964), In the past 25 years many nature lovers in the state have made the pilgrimage to Lime Pond attempting to rediscover Calypso in New Hampshire. To date, the site has not yielded the beautiful orchid again and the seekers return from their expeditions with only white lime-encrusted boots and scores of black-fly bites. The brightly-colored flower and position of the column suggest that Calypso is pollinated by bees. Although the orchid produces no nectar it does display the yellow bristles that are believed to mimic anthers. The European plants are reported to have a vanilla-like scent which attract bees who feed on the blueberry (Vaccinium) flowers that bloom at the same time (S. Nilsson, 1979). Mosquin (1970) has studied the reproductive biology of Calypso bulbosa in Alberta, Canada, and has identified bumblebees of the genera Bom- bus and Psithyrus (Apidae) as carrying pollinia of Calypso on their thoraxes. Stoutamire (1971) listed the bumblebees Bombus terna- rius and B. vagans (Apidae) as pollinators of Calypso. Unlike the apparent situation in northwestern North America, few plants in the northeast form fruit, a factor which may account for the rarity of the species here. Nylander (1922) observed birds attacking the flowers of Calypso in Woodland, Maine, a practice that he believed accounted for the low percentage of seed-set in this species. When the flower is not pollinated it remains fresh for about three weeks (Nylander, 1935; Mosquin, 1970). Calypso propagates vegetatively through the production of coralline rhizomes (Mously, 1925). Correll (1950) reported that the Indians of British Columbia eat the corms of Calypso which contain a large amount of a mucilagin- ous substance. Every year during my orchid hunting forays in New Hampshire, I discover new locations with habitats suitable for Calypso and faith- 1985] Brackley — Orchids of New Hampshire 97 fully return the following May to seek the elusive plant. That I have not yet met with success is not surprising when one considers that it was only after 25 years of searching that Calypso revealed itself to Morris and Eames (1929). The thought of finding Calypso lingers long after its blooming dates have passed and the lines from Frost’s (1964) poem “The Encounter” keep returning to my mind. “Sometimes | wander out of beaten ways Half looking for the orchid Calypso” GENERAL DISTRIBUTION: Circumboreal, extending southward in the Rocky Mountains to Arizona and California. 17. Corallorhiza [Hall.] Chatelain, Spec. Inaug. de Corallorhiza 6. 1760. A small genus of eight to ten species which have a center of distribution in Mexico (Leur, 1975), the name for the genus was already well established when Haller used it in his pre-Linnean work published in 1742. The author of Cora/lorhiza must be attri- buted to Chatelain who published the first description of this genus after 1753. Corallorhiza is from two Greek words signifying “coral-root.” Although the members of this genus do not actually have roots they do possess an intricately branched underground rhizome that is coralline in appearance. Corrallorhizas have little or no chlorophyll and are saprophytes associated with mycorrhizal fungi and/ or indi- rect parasites on the roots of living trees. Two species of Corallorhiza commonly occur in New Hampshire and there is a historic record for a third, Corallorhiza odontorhiza. Key to the common species of Coral/lorhiza in New Hampshire 1. Plant yellowish, usually less than 20 cm tall; lip 3-5 mm long; blossoming in spring or early summer; plants of wet areas.... SERENA RTE OMAR ES ona a eee ae ee 1. Corallorhiza trifida 1. Plant purplish, usually greater than 20 cm tall; lip 5-9 mm long; usually blossoming in late summer; plants of upland woods oO ee eee ee eee re eee 2. Corallorhiza maculata I. Corallorhiza trifida Chat. Spec. Inaug. de Corallorhiza 8. 1760. 98 Rhodora [Vol. 87 Linnaeus (1753) originally described this species as Ophrys coral- lorhiza, and it has also been known as Corallorhiza innata. Coral- lorhiza trifida is circumpolar in distribution with the Eurasian and northernmost North American plants referable to var. trifida. Vari- ety verna (Nutt.) Fern. occurs in our area and is distinguished by a lighter, more yellowish color and a white, unspotted, abruptly- tipped lip. This difference was sufficient to lead Nuttall to consider it as a separate species, which he called C. verna. The specific epithet trifida is from Latin and means “split into three,” referring to the three-lobed lip. The varietal name verna means “spring,” the season of flowering. COMMON NAME: Early Coral-root FLOWERING DATES IN NEW HAMPSHIRE: May through June. DESCRIPTION: Plant froma fragile much-branched rhizome, erect, yellowish, glabrous, averaging approximately 18 cm in height, with 2-3 tubular sheaths enclosing the stem. Inflorescence a loosely flow- ered raceme of few-20 flowers. Floral bracts minute, acute. Flow- ers spreading yellowish-white on short slender pedicels. Sepals lanceolate, concave. Lateral petals linear-lanceolate. Lip white, unspotted or with very few basal spots, obovate, 3-lobed. Lateral lobes of lip near base, short, triangular. Mid-lobe expanded, bluntly tipped. Column curved, slightly clavate. Capsule obovoid, pendant. CHROMOSOME NUMBER: 2n = 42. One of our earliest orchids to blossom, the flowers of Corallorhiza trifida var. verna are usually in evidence by May. The yellowish to yellowish-green color of the plant signifies that this taxon does possess a small amount of chlorophyll, unlike the majority of spe- cies in the genus. The characteristic coralline rhizomes which lie beneath the surface of the forest floor are much-branched and knobby in appearance. On the surface of the rhizome of the Early Coral-root, Thomas (1893) found tiny papillae with protruding tufts of hairs. Campbell (1970) reported that various fungi, with white or yellow hyphae, enter the rhizome through the protruding hairs and advance into the cortex. Her study also revealed that some of the fungal hyphae found in the Early Coral-root are addi- tionally attached to the roots of Cedar (Thuja) or Spruce (Picea) trees. The orchid presumably obtains some of its nourishment from 1985] Brackley — Orchids of New Hampshire 99 the trees and therefore functions as an indirect parasite through the fungal intermediaries. In order to see the Early Coral-root in blossom one must venture into a wet coniferous or mixed forest at the height of Black-fly season. Since this species is shade tolerant it can thrive in the dark- ness that characterizes our Cedar woods. In more southern areas Corallorhiza trifida var. verna is often found growing along the edges of forest streams. The Early Coral-root can occasionally be found in bogs. My first meeting with this orchid was in a bog in Merrimack County where it was found growing tucked beneath the shrubs at the edge of the bog mat. Wherry (1927) reported that the apparent optimum soil for this species is one with a minimum-acid reaction, and Stuckey (1967) reported this species growing in soils with a pH of 5.4 in Rhode Island. A characteristic of this genus is the joined bases of the lateral sepals, which are somewhat fused with the lip forming a small sac-like spur (Bingham, 1939). It has not been determined if nectar is produced in the Early Coral-root. On the upper surface of the lip are two longitudinal ridges that probably serve to direct insects to the spur. Corallorhizas are protandrous. The anther produces two pollen-masses which remain under the anther lid. At this time the column renders the opening of the blossom too narrow for insects to reach the spur. When an insect probes the flower, it trips open the anther lid causing the pollen-mass to become attached to its head. When the stigma is mature the flowers open widely and the next insect visitor with attached pollen-masses easily transfers them to the sticky stigma. In Europe, Silen (1906) reported that the Hover-fly Syrphus cinctellus (Syrphidae) visited Corallorhiza trif- ida. Catling (1980a) suggested that this species may be partly auto- gamous. | have found the beetle Anthobium convexium (Sta- phylinidae) on the flowers of the Early Coral-root but they appeared to be only feeding on the pollen-masses. When fertilization occurs in the Early Coral-root the ovaries increase in size and turn a greenish color, possibly indicating that most of the chlorophyll in the plant is concentrated in this organ. Corallorhiza trifida var. verna is a good seed producer. Propaga- tion in this species is primarily by sexual means, as vegetative mul- tiplication in the Early Coral-root is reportedly not as effective as in most orchids (Summerhayes, 1951). 100 Rhodora [Vol. 87 GENERAL DISTRIBUTION: Newfoundland west through the Great Lakes area to British Columbia and Oregon, south to Pennsylvania. 2. Corallorhiza maculata Raf., Amer. Monthly Mag. and Crit. Rev. 2: 119. 1817. This new world species of Corallorhiza was originally described by Rafinesque from plants found in New York. The species has also been known as C. multiflora. The epithet maculata is Latin for “spotted,” describing the purple spots on the lip. COMMON NAMES: Spotted Coral-root, Large Coral-root FLOWERING DATES IN NEW HAMPSHIRE: late July through early September, rarely earlier. DESCRIPTION: (Anextremely variable species). Plant from a coral- line rhizome, erect, glabrous, succulent, purplish, with sheaths enclosing the stem. Raceme few- to many-flowered. Floral bracts obscure. Flowers spreading, on stout pedicels. Sepals purplish, oblong-lanceolate. Lateral sepals joined under the lip forming a small spur-like sac. Lateral petals slightly smaller and of a lighter purplish color than the sepals, oblong-lanceolate. Lip three-lobed, white with magenta spots and 2 longitudinal median ridges. Lateral lobes small, inrolled, directed forward. Mid-lobe expanded, broadly rounded. Column yellow, spotted with magenta, curved. Capsule ellipsoid, pendent. CHROMOSOME NUMBER: 2n = 42 (Moore, 1973). The Spotted Coral-root is one of the most widespread of all the orchids of our flora. Corallorhiza maculata \acks chlorophyll entirely, a feature that led earlier writers to label the species “degen- erate” (Gibson, 1905). But this species is rich in other color pig- ments. A typical plant usually has a purplish stem, yellowish to purplish sepals and lateral petals, and a white lip spotted with magenta. Overall, C. maculata is a handsome plant. Several different color types of Corallorhiza maculata exist, some of which have been described as varieties (Bartlett, 1922). These are characterized as follows: |. var. flavida (Peck) Farwell: this taxon lacks the purple antho- cyanins which usually color the stems and sepals, resulting ina yellow plant with a white unspotted lip. 1985] Brackley — Orchids of New Hampshire 101 2. var. intermedia Farwell: an overall brownish plant, perhaps representing only older specimens of var. maculata (Case, 1964). 3. var. punicea H. H. Bartlet: a more slender version with trans- parent sheaths and reddish-purple stems. The color differences in this species are noticeable in the field but, upon drying, all plants of C. macu/ara turn a uniform reddish-brown. Usually it is late summer before the Spotted Coral-root emerges and opens its flowers. Plants of Cora/llorhiza maculata can be found growing singly or in clumps. In New Hampshire the favored habitat for this species is in Beech-Maple-Hemlock forests. At that time of year other saprophytes indigenous to this community are also apparent. Indian Pipes (Monotropa uniflora) and Beech-drops (Epifagus virginiana) are usually associated with the Spotted Coral- root. It is easy to confuse the Spotted Coral-root with Beech-drops when the plants are very young. If one remembers that Beech-drops exhibit a branching habit, whereas the Spotted Coral-root never branches, the two plants can be readily distinguished. In 1980, when photographing the early-blooming Corallorhiza trifida var. verna in a wet streamside habitat, I noticed a clump of pale yellow succulent-looking stalks pushing up through the leaf litter. When I returned to the station two weeks later, I was greeted by the sight of a robust colony of Corallorhiza that was of such an unusual form, and flowering at so early a date, that I was certain | had discovered something special. The plant, however, would key out only to C. maculata. The entire plant was pale yellow but, unlike the description of var. flavida, the white lips displayed magenta spots. Never before had I encounted this species flowering so early or living in such a habitat. Later, I read of Morris and Eames (1929) being shocked to find the yellow form of the Spotted Coral-root in bloom as early as June 13th. This is truly a highly diversified species. In her study of the fungal associations of Corallorhiza in Michi- gan, Campbell (1970) found rhizomorphs of the Honey Mushroom (Armillariella mellea) associated with the hairs of papillae on the Spotted Coral-root rhizomes. The same rhizomorphs were also found penetrating roots of adjacent trees. It is interesting that the fungal associates of C. maculata are quite different from those found in C. trifida var. verna (Campbell 1970). 102 Rhodora [Vol. 87 The Spotted Coral-root appears to be morphologically adapted for insect pollination, but I have not found any mention of pollina- tors in the literature for this species. Catling (1980a), however, reported that evidence indicates this orchid may be partly auto- gamous. There is an obvious need for further investigation into the reproductive biology of this taxon. GENERAL DISTRIBUTION: Newfoundland south to North Carolina, west through the Great Lakes area to Washington, and south through the mountains to Arizona; Mexico and Guatemala. ACKNOWLEDGMENTS I thank Garrett E. Crow, A. Linn Bogle, Owen M. Rogers and Leslie A. Garay for their helpful critisms of the manuscript. I am grateful to all the herbarium curators who gave their time and assistance to me. Many thanks are extended to John Burger and Wally Morse for identifying the insects collected in this study. My deepest appreciation is extended to J. Korpi, C. Levesque, K. Price, C. Bartholomew and F. Smith for their help and moral sup- port. For introducing me to the study of Botany, | thank David Boufford and David Gregory. Finally, | wish to thank the people of New Hampshire for sharing their knowledge of the flora and for their many kindnesses. This work was supported in part by a grant from the American Orchid Society and by two UNH Central University Research Grants (CURF S135 and S167). 1985] Brackley — Orchids of New Hampshire 103 DISTRIBUTION MAPS OF 43 NEW HAMPSHIRE ORCHIDS Bold-face numbers refer to genus in the key and text; other number is species under that particular genus. Each dot on the map represents an historic or extant record(s) for the species within that town or township. Maps have been updated to reflect field work done since 1981. 1-3 CYPRIPEDIUM PUBESCENS 1-4 CYPRIPEDIUM PARVIFLORUM 104 Rhodora [Vol. 87 3-1 COELOGLOSSUM VIRIDE ssp. BRACTEATUM 1985] Brackley — Orchids of New Hampshire 105 4-6 PLATANTHERA HOOKERI i 4-7 PLATANTHERA OBTUSATA 4-8 PLATANTHERA CLAVELLATA ssp. OBTUSATA 106 Rhodora [Vol. 87 1 a 5 @ >4. \ ro | . As “) ies cf 4 ba: Diane 3% “ 5-1 EPIPACTIS HELLEBORINE 1985] Brackley — Orchids of New Hampshire 107 ty Z 6-3 LISTERA AURICULATA 7A. ae 28, ares wt F a7 t X ANS AL EES f waas: 9-1 SPIRANTHES LUCIDA 108 Rhodora [Vol. 87 , a = : 9-2 SPIRANTHES ROMANZOFFIANA 9-3 SPIRANTHES CERNUA 9-5 SPIRANTHES CASEI LP KS , 7 x re ot f le ™s Ye 10-1 ISOTRIA VERTICILLATA 10-2 ISOTRIA MEDEOLOIDES 1985] Brackley — Orchids of New Hampshire 109 14-2 MALAXIS UNIFOLIA 15-1 LIPARIS LOESELII 110 Rhodora [Vol. 87 J6 By : a4 re, AY AA Ly rae my rf A ‘g P a 7 — miu ~ ° Se a a “ oa) . WM noe ey; A YR - 17-1 CORALLORHIZA TRIFIDA 17-2 CORALLORHIZA MACULATA 1985] Brackley — Orchids of New Hampshire 111 LITERATURE CITED ACKERMAN, J.D. 1975. Reproductive biology of Goodvera oblongifolia (Orchi- daceae). Madrofio 23: 191-198. AND M.R. MEsSLER. 1979. Pollination biology of Listera cordata (Orchi- daceae). Amer. J. Bot. 66: 820-824. Ames, O. 1906. Habenaria orbiculata and H. macrophylla. Rhodora 4: |-S. 1921. 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Bull. Torrey Bot. Club 6: 329-330. ARTHUR, A. P. 1962. Adults of the European skipper Thymelicus lineola (Ochs.). Proc. Entomol. Soc. Ontario 92: 190-191. Auctair, A.N. 1972. Comparative ecology of the orchids Ap/ectum hymale and Orchis spectabilis. Bull. Torrey Bot. Club 99: I-10. AYENSU, E.S. AND R. A. DEFILIPPS. 1978. Endangered and Threatened Plants of the United States. Smithsonian Institution and World Wildlife Fund, Inc., Washington, DC. BaLDWIN, H. 1894. The Orchids of New England. John Wiley and Sons, New York. BALDWIN J. T. AND B. M. SpegEsE. 1957. Chromosomes of Pogonia and of its allies in the range of Gray’s Manual. Amer. J. Bot. 44: 651-653. BARKER, E. E. 1931. You also can grow orchids as I do in N. Y. State. Better Homes and Gardens. April issue. BARTLETT, H. H. 1922. Color types of Corallorrhiza maculata Raf. Rhodora 24: 145-148. BATCHELDER, F. W. 1900. Flora of Manchester and vicinity. Proc. Manchester Institute of Arts and Sciences |: 1-158. 1909. Preliminary list of plants growing without cultivation in the vicin- ity of Manchester, New Hampshire. Proc. Manchester Institute of Arts and Sciences. 4: 3-59. BiGELow, J. 1824. Florula Bostoniensis. 2nd ed. Cummings, Hilliard, and Co., Boston. 112 Rhodora [Vol. 87 BINGHAM, M.T. 1939. Orchids of Michigan. Bull. Cranbrook Inst. Sci. 15: 1-87. BRACKLEY, F. E. 1979, Cypripedium reginae rediscovered in New Hampshire. Rhodora 81: 579-580. BRITTON, N. L. 1901. Manual of the Flora of the Northern States and Canada. New York. CAMPBELL, E.O. 1970. Morphology of the fungal association in three species of Corallorhiza in Michigan. Mich. Bot. 9: 108-113. Cask, F. W. JR. 1964a. Orchids of Western Great Lakes Region. Cranbrook Institute, Bloomfield Hills, Michigan. 1964b. A hybrid twayblade and its rarer parent, Listera auriculata, in northern Michigan. Michigan Bot. 3: 67-70. ——. 1971. Isotria medeoloides, the smaller Whorled Pogonia, in Michigan. Michigan Bot. 10: 39-43, CATLING, P.M. 1978. Taxonomic notes on Spiranthes casei Catling and Cruise and Spiranthes Xintermedia Ames. Rhodora 80: 377-389. 1980a. Autogamy in north-eastern North American orchids. Bot. Soc. Amer. Misc. Ser. 158: 20. 1980b. Rain-assisted autogamy in Liparis loesellii (L.) L. C. Rich (Orchidaceae). Bull. Torrey Bot. Club 107: 525-529. CATLING, P. M. AND J. E. Cruise. 1974. Spiranthes casei, a new species from northeastern North America. Rhodora 76: 526-536. Copy, W. J. AND D. Monro. 1980. The genus Listera (Twayblades) in New Brunswick. Canad. Field-Nat. 94: 443-446, Corrett, D. S. 1941. A new Spiranthes hybrid from New Hampshire. Amer. Orchid Soc. Bull. 9: 241. 1950. Native Orchids of North America. Stanford Univ. Press, Stanford, CA. CoUNTRYMAN, W. D. 1978. Rare and endangered vascular plant species in Ver- mont. New England Botanical Club in cooperation with the U. S. Fish and Wildlife Service, Region 5 [Newton Corner, MA]. Crow, G. E. er al. 1981. 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THE NATURE CONSERVANCY NEW HAMPSHIRE NATURAL HERITAGE PROGRAM 54 PORTSMOUTH STREET CONCORD, N.H. 03301 AN AWARD FOR THE SUPPORT OF BOTANICAL RESEARCH IN NEW ENGLAND, U.S.A. The New England Botanical Club is offering an award of $1,000 in support of botanical research to be conducted in the New England region during 1985. It is being made to stimulate and encourage botanical research on the New England flora and to make possible visits to the New England region by those who would not otherwise be able to do so. The award will be given to the graduate student submitting the best research proposals dealing with field studies in systematic botany and plant ecology, but proposals for research in other areas of botany will also be considered. This award is not limited to graduate students at New England institutions. The NEBC’s support must be acknowledged in any publications resulting from this study. It is encouraged that papers based on this research be submitted to RHODORA, the Club’s journal, for possible publication—subject to standard review processes. The New England Botanical Club hopes to be able to make this award on an annual basis. Applicants should submit a proposal of no more than three double spaced pages, including a budget (the budget will not affect the amount of the award), and their Curriculum Vitae. Two letters, one from the student’s major professor, in support of the proposed research are also required. Proposals and supporting letters should be sent before 28 February 1985 to: Awards Committee, The New England Botanical Club, 22 Divinity Avenue, Cambridge, MA 02138. The recipient of the award will be notified by 30 April 1985. 118 ASPT HERBARIUM TRAVEL AWARDS The American Society of Plant Taxonomists is pleased to announce the availability of competitive awards for travel by grad- uate students to the nation’s herbaria. The awards will not exceed $500 and will be used to help pay expenses to and from any herba- rium (or herbaria) in the United States and per diem expenses during the visit. Competitions for awards will be held twice a year: The first competition deadline is | January 1985, with the second deadline | July 1985. The grants program will last a minimum of three years (six competitions). Interested Master’s or Ph.D. gradu- ate students should send a curriculum vitae, two letters of recom- mendation (including one from the major professor), a two or three page outline of the proposed research emphasizing the role that the visit to the herbarium will play, and a letter from the Head Curator, Chairman or Director of the institution(s) to be visited indicating willingness to receive the visitor. Awards will be announced by | March from the January competition and during the annual ban- quet of the ASPT from the July competition. Students are encour- aged to obtain additional funds from their home institutions (or elsewhere) to extend their research visits even further. This competi- tion is open to students of both cryptogamic and phanerogamic groups. Completed applications and additional questions should be directed to Tod F. Stuessy, Chairman, ASPT Committee for Sys- tematics Collections, Department of Botany, Ohio State University, 1735 Neil Avenue, Columbus, OH 43210. Phone: (614) 422-5200 or (614) 422-8952. Vol 86, No. 848, including pages 397-554, was issued December 20, 1984 119 INSTRUCTIONS TO CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simultaneously elsewhere, either in whole or in part. Manuscripts should be submitted in triplicate (an original and two xerox copies) and must be double-spaced (at least 3/8 of an inch) throughout including footnotes, figure legends, and refer- ences. Please do not use corrasable bond. The list of legends for figures and maps should be provided on a separate page. Footnotes should be used sparingly. Do not indicate the style of type through the use of capitals or underscoring, particularly in the citation of specimens. Names of genera and species may be underlined to indi- cate italics in discussions. Specimens citations should be selected critically, especially for common species of broad distributions. Sys- tematic revisions and similar papers should be prepared in the for- mat of “A Monograph of the Genus Malvastrum”, S. R. Hill, Rhodora 84: 1-83, 159-264, 317-409, 1982, particularly with refer- ence to indentation of keys and synonyms. Designation of a new taxon should carry a Latin diagnosis (rather than a full Latin de- scription), which sets forth succinctly just how the new taxon is distinguished from its congeners. Papers of a floristic nature should follow, as far as possible, the format of “Annotated list of the ferns and fern allies of Arkansas”, W. Carl Taylor and Delzie Demaree, Rhodora 81: 503-548, 1979. For bibliographic citations, refer to the - Botanico-Periodicum-Huntianum (B-P-H, 1968), which provides standardized abbreviations for journals originating before 1966. All abbreviations in the text should be followed by a period, except those for standard units of measure and direction (compass points). For standard abbreviations and for guidance in other matters of biological writing style, consult the CBE Style Manual, Sth ed. (original title: Style Manual for Biological Journals). In preparing figures (maps, charts, drawings, photos, etc.) please remember that the printed plate will be 4 X 6 inches; be sure that your illustrations are proportioned to reduce correctly, and indicate by blue pencil the intended limits of the figures. (Some “turn-page” figures with brief legends will be 3 1/2 X 6 in.) Magnification/ reduction values given in text or figure legends should be calculated to reflect the actual printed size. An Abstract and a list of Key Words should be sup- plied at the beginning of each paper submitted, except for a very short article or note. RHODORA January 1985 CONTENTS The orchids of New Hampshire Frances E. Brackley NEBC Research Award Notice ASPT Herbarium Travel Awards Notice Instructions to Contributors Vol. 87, No. 849 119 . inside back cover JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB eave aad AT: . soe 4 ic 2 Foy * so we a: a o =*s +" » > ae . ° . =? 5 ee 7. oe si iat ’ hy pare ante a e 7? April 1985 No. 850 Vol. 87 Che New England Botanial Club, Ine. Botanical Museum, Oxford Street, Cambridge, Massachusetts 02138 Conducted and published for the Club, by NORTON H. NICKERSON, Editor-in-Chief DAVID S. BARRINGTON RICHARD A. FRALICK A. LINN BOGLE GERALD J. GASTONY WILLIAM D. COUNTRYMAN MICHAEL W. LEFOR GARRETT E. CROW NORTON G. MILLER ROBERT T. WILCE RHODORA.—Published four times a year, in January, April, July, and October. A quarterly journal of botany, devoted primarily to the flora of North America. Price $20.00 per year, net, postpaid, in funds payable at par in the United States currency at Boston. Some back volumes and single copies are available. Information and prices will be furnished upon request. Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to RHODORA, Botanical Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive the next number of RHODORA, changes of address must be received prior to the first day of January, April, July or October. Scientific papers and notes relating to the plants of North America and floristically related areas will be considered by the editorial committee for publication. Articles concerned with systematic botany and cytotaxon- omy in their broader implications are equally acceptable. Brevity is urged whenever possible in all papers. Short items will be published on otherwise blank end pages as soon as possible, even if they appear ahead of longer articles already accepted. All manuscripts should be submitted in TRIPLICATE AND MUST BE DOUBLE (AT LEAST 3/8 OF AN INCH) OR TRIPLE- SPACED THROUGHOUT. Please conform to the style of recent issues of the journal. See “Instructions to Contributors to RHODORA” at the end of each issue. Extracted reprints, if ordered in advance, will be furnished at cost. RHODORA assesses modest page charges. Address manuscripts and proofs to: Joan Y. Nickerson Managing Editor, RHODORA Phippen-LaCroix Herbarium, Dept. of Biology Tufts University Medford, Mass. 02155 Second Class Postage Paid at Boston, Mass. PRINTED BY THE LEXINGTON PRESS, INC LEXINGTON, MASSACHUSETTS Cover illustration Cypripedium reginae Walt., the Showy Lady’s-slipper, was presumed until recently to be extirpated from New Hampshire; five sites for this exquisite orchid are now known. Original artwork by Tess Feltes, Illustrator. Rhodora (ISSN 0035-4902 JOURNAL OF ae NEW ENGLAND BOTANICAL CLUB Vol. 87 April 1985 No. 850 SYSTEMATICS OF TALINUM PARVIFLORUM NUTT. AND THE- ORIGIN: OF 7. TERETIFOLIUM PURSH (PORTULACACEAE) M. ELoise BROWN CARTER AND WILLIAM H. Murby ABSTRACT Talinum appalachianum Wolf, known from only one station in Alabama, is not distinct from T. parviflorum Nutt. Plants of 7. parviflorum were sampled from Texas, Arkansas and Kansas and were compared with 7. appalachianum. Plants of the four populations had the same chromosome number (2n = 24), very late diurnal anthesis, and were predominantly self-pollinating. Population means for four floral traits overlapped: petal length. number of stamens, style length and number of ovules. Talinum teretifolium Pursh appears to be an allotetraploid derivative of T. mengesii Wolf and 7. parviflorum. Crosses between the latter two species produced sterile hybrids similar to 7. reretifolium in floral morphology and pattern of diurnal anthesis. Fertile polyploid flowers, which developed after colchicine treatment, pro- duced fertile offspring when crossed to 7. reretifolium, selfed or sibbed. Key Words: Jalinum, allopolyploidy, hybridization, rock outcrop, southeastern U.S. INTRODUCTION Talinum (Portulacaceae) has approximately 50 species distrib- uted in Africa, Asia and North and South America. Wilson (1932) described 31 species from North America. Eight species of Ta/inum have been reported for the southeastern United States (Bogle, 1969). Talinum paniculatum (Jacq.) Gaertn. and T. triangulare (Jacq.) Willd. are flat-leaved, horticultural escapes with worldwide distribu- tions. Six species have terete leaves and inhabit shallow soils of rock outcrops. Talinum parviflorum Nutt., 2n = 48 (Steiner, 1944) and 2n = 24 (Hornberger & Smith, pers. comm.), and 7. calycinum Engelm., 2n = 24 (Steiner, 1944), range widely west of the Missis- 121 122 Rhodora [Vol. 87 sippi River with a few populations in southern Illinois (Mohlen- brock, 1975). Distribution ranges for the remaining four species are wholly east of the Mississippi River: 7. reretifolium Pursh, 2n = 48 (Steiner, 1944; Murdy, 1968; Murdy, Johnson, & Wright, 1970), occurs on granite, sandstone and serpentine from Georgia to Penn- sylvania; 7. mengesii Wolf, 2n = 24, 48 (Steiner 1944; Murdy et al., 1970), occurs on sandstone and granite in Georgia, Alabama, and Tennessee; 7. appalachianum Wolf, 2n = 24 (Steiner, 1944), con- sists of a few populations within a one-mile radius on gneiss in central Alabama, and 7. calcaricum Ware, 2n = 48 (Krebs, 1971, Master’s thesis, Emory Univ., GA), is restricted to limestone in central Tennessee and northwestern Alabama (Figure 1). The six terete-leaved species have short stems bearing numerous succulent leaves and produce flowers on long, slender, cymosely- branched scapes. All produce ephemeral flowers which open in the afternoon between 1200 hours and 1800 hours EDT and close by 2000 hours EDT. The species are distinguished primarily by floral characteristics including length of petals, number of stamens, length of style relative to stamens, petal color, persistence of sepals, and seed color (Ware, 1967; Bogle, 1969; Black & Murdy, 1972). T. mengesii T. teretifolium T. parviflorum T. appalachianum Figure |. Distribution of Tal/inum appalachianum, T. parviflorum, T. mengesii, and T. teretifolium. 1985] Carter & Murdy — Talinum 123 The research reported in this paper tested two hypotheses: 1) Talinum appalachianum is not a species distinct from 7. parviflo- rum and 2) T. teretifolium is an allopolyploid derivative of 7. men- gesti and T. parviflorum. In 1937, R. M. Harper discovered an isolated population of Tali- num on rocks in central Alabama. Wolf (1939) noted a resemblance to T. parviflorum but named it 7. appalachianum. The species, 7. appalachianum, was not challenged by later investigators (Steiner, 1944; Bogle, 1969; Black & Murdy, 1972) in part because the nearest reported population of 7. parviflorum occurred hundreds of miles west of this isolated population, and because Steiner (1944) had reported a chromosome number of 2n = 48 for a population of T. parviflorum in Arkansas and 2n = 24 for 7. appalachianum. The species were thought to be at different ploidal levels until Horn- berger and Smith (pers. comm.) reported 2n = 24 for a T. parviflo- rum population from Washington County, Arkansas. Based on the evidence from comparative morphology, distribu- tion patterns and chromosome numbers, Murdy (1968) suggested that Talinum teretifolium was an allotetraploid and that 7. mengesii had been one of its ancestral parents. Cytological analysis of inter- ploidal hybrids between the two species supported the hypothesis, and the use of Anderson’s method of extrapolated correlates (An- derson, 1949) suggested that a diploid species similar to T. appalachi- anum could have been the other ancestral parent (Black & Murdy, 1972). MATERIALS AND METHODS Talinum mengesii populations (1E5) were collected from one county in Alabama and four counties in eastern to central Georgia. Talinum parviflorum populations (6E8) were collected from single sites west of the Mississippi River in Arkansas, Kansas and Texas, and one population of T. appalachianum (9) was sampled from its only known station in Alabama. Talinum teretifolium populations (10E14) were sampled from five counties across Georgia. Locations for each population are given in Table 1. Fourteen to twenty individuals were sampled in the field either from seed or rhizome. To minimize selection of the same genotype and yet provide a representative sample, one individual or capsule was collected per patch of plants. A patch was defined as a group of 124 Rhodora [Vol. 87 Table |. Sample site locations for populations of Talinum Population Number Location T. mengesii l Blake’s Ferry, Randolph County, Alabama 2 Highway 34, Franklin, Heard County, Georgia 3 Powers Ferry Crossing, Coweeta County, Georgia 4 Costley Bridge Road, Rockdale County, Georgia 5 Highway 138, Walton County, Georgia T. parviflorum 6 Devil’s Den State Park, Washington County, Arkansas 7 Sedan, Chautauqua County, Kansas 8 Inks Lake State Park, Burnet County, Texas T. appalachianum 9 Highway 22 and Coosa River, Coosa County, Alabama T. teretifolium 10 Carl, Barrow County, Georgia I] Pendergrass, Jackson County, Georgia 12 Echols Mill, Oglethorpe County, Georgia 13 North High Shoals, Oconee County, Georgia 14 Kent Rock Road, Walton County, Georgia plants separated from the next group by at least one meter. Plants subsequently grown from seed or transplants were potted in a mix- ture of granite outcrop soil, sand and peat (1:1:1) and were main- tained in the greenhouse. One flower from each plant was collected and preserved in FAA for measurement of floral characters. Lengths of petals, pistils and styles were measured to the nearest 0.5 mm; number of stamens and number of ovules per ovary were recorded for each flower. Vouchers were deposited at GEO. Chromosome counts were determined for three populations of Talinum parviflorum and one population of 7. appalachianum. Plants were stripped of roots, and the rhizomes were placed in moist paper toweling. New roots that developed within 4 days were excised from the rhizome, placed in a 0.2% colchicine solution for 3 hours, and then transferred to Carnoy’s fixative (3 absolute alcohol : I acetic acid) where they remained for at least 24 hours. Fixed root tips were hydrolyzed in 1 N HCI and acetocarmine squash prepara- tions were made. Chromosomes were counted for a minimum of three plants to establish the chromosome number for a particular population. 1985] Carter & Murdy — Talinum 125 Plants were destaminated in bud and subsequently hand polli- nated in an isolation chamber. Mature seeds were collected and placed in dry storage for 5 weeks, refrigerated at 4OC in a moist petri-dish for a minimum of 6 weeks and germinated in sterilized granite outcrop soil amended with sand (1:1). Forty-nine crosses were made using 13 different Ta/linum appalachianum plants as female parents and pollen from several 7. parviflorum individuals. Seeds from these crosses were pretreated and germinated. The resulting plants were grown to maturity and allowed to self- pollinate. Crosses (107) were attempted using 17 different female parents of Talinum appalachianum and a mixture of pollen from several 7. mengesii individuals. Seeds from these crosses were pretreated, germinated and grown to maturity in the greenhouse. Colchicine (1% in lanolin) was used to induce polyploidy in 15 of the sterile, interspecific hybrids. The branches that developed from treated nodes bore fertile flowers and were, therefore, assumed to be poly- ploid. Crosses were made between these flowers and those of the tetraploid, Talinum teretifolium. Five flowers from each hybrid plant were collected and preserved in FAA for floral character anal- ysis. Pollen fertility was determined by staining the pollen with iodine and dividing the number of normal sized, starch-filled grains by the total number. Fruit set (percent of pollinations resulting in capsules) was determined from self and sib crosses of fertile flowers. Seed set (number of mature seeds per capsule) was recorded for successful crosses. Branches from hybrid plants are deposited at GEO. RESULTS One population of Talinum appalachianum from Alabama and three populations of 7. parviflorum from Arkansas, Texas and Kansas had chromosome counts of 2n = 24. The four populations could not be distinguished on the basis of vegetative habit and they were similar in floral morphology. Talinum appalachianum had slightly larger petals than 7. parviflorum, but population means overlapped for number of stamens, style length and number of ovules (Table 2). In all four populations, anthesis occurred very late in the day between 1630 and 1800 hours EDT, and the flowers were self-pollinating at flower closure around 2000 hours EDT. 126 Rhodora [Vol. 87 Table 2. Floral traits for Ta/inum populationsp No. Petal No. Style No. Population Plants Length Stamens Length Ovules T. mengesii I 20 8.48 (0.53) 52.30 (9.15) 4.04 (0.30) 35.05 (5.92) 2 20 8.96 (0.68) 61.55 (9.11) 4.64 (0.49) 35.80 (8.69) 3 20 8.61 (0.45) 60.20 (7.52) 4.23 (0.42) 39.90 (6.03) 4 20 10.78 (0.73) 67.35 (6.98) 6.20 (0.35) 46.75 (8.27) 5 20 10.00 (0.61) 71.20 (7.49) 5.81 (0.44) 59.50 (5.40) Species Mean 9.37 (0.99) 62.52 (7.24) 4.98 (0.97) 43.40 (10.13) T. parviflorum 6 16 4.70 (0.49) 6.69 (1.57) 1.08 (0.19) 20.38 (5.69) 7 11 4.82 (0.48) 7.09 (1.88) 1.27 (0.23) 28.27 (6.88) 8 15 3.97 (0.31) 4.87 (0.34) 0.68 (0.17) 23.95 (3.40) T. appalachianum 9 18 5.17 (0.42) 5.72 (0.87) 1.12 (0.17) 24.78 (3.92) Species Mean 4.66 (0.50) 6.09 (1.00) 1.04 (0.25) 24.35 (3.24) T. teretifolium 10 20 7.08 (0.59) 17.40 (3.44) 2.56 (0.26) 37.65 (5.67) II 20 6.88 (0.65) 20.95 (4.74) 2.96 (0.19) 48.75 (6.16) 12 20 6.46 (0.53) 18.20 (2.86) 2.66 (0.23) 38.75 (6.10) 13 20 6.10 (0.45) 15.90 (1.21) 2.83 (0.23) 31.50 (4.16) 14 20 6.73 (0.44) 17.30 (2.81) 2.76 (0.21) 41.65 (6.28) Species Mean 6.65 (0.38) 17.95 (1.87) 2.75 (0.15) 39.66 (6.29) T. appalachianum X T. mengesii (synthetic hybrid) 15 7 8.67(0.29) 25.80 (2.42) 3.14 (0.14) 48.23 (1.77) 16 8 8.49 (0.21) 24.45 (3.02) 2.97 (0.28) 47.13 (7.82) p Population means with standard deviations in parentheses. Crosses between Talinum appalachianum and T. parviflorum resulted in 55% fruit set with mean seed set of 22.7 seeds per capsule (n= 49) as compared to 23.1 seeds per capsule from intrapopulation control pollinations of 7. appalachianum (n = 10). Ten hybrid plants from crosses of 7. appalachianum with T. parviflorum, Texas, and eight hybrid plants from crosses of 7. appalachianum with 7. parviflorum, Arkansas, were grown to maturity. All hybrid plants bore fertile flowers which ranged in pollen fertility from 68% to 85%, and which self-pollinated to produce fruit with mean seed set of 9 seeds per capsule. Ten 7. parviflorum plants grown under the same environmental regime had 77% to 92% pollen fertility and 1985] Carter & Murdy — Talinum 127 produced fruit with an average of 11.0 seeds per capsule. Hybrids resulting from crosses with 7. parviflorum, Kansas, did not flower during the first growing season. An attempt to reconstruct the origin of the tetraploid species, Talinum teretifolium, using its presumed progenitors, 7. mengesii and T. appalachianum (to be referred to hereafter as 7. parviflorum, Alabama), was initiated in 1981. Only 6.5% of the 107 crosses attempted between 7. mengesii and T. parviflorum, Alabama, resulted in fertile capsules with a mean of 4.5 seeds per capsule. Germination was attempted with 17 seeds; 15 germinated and sur- vived. All 15 hybrid plants which were grown to maturity in the greenhouse were sterile; mean pollen fertility was less than 15% and fruit set was zero. Floral morphology and time of diurnal flower opening for the hybrids fell within the range of variability for T. teretifolium. They had a higher value for petal length, stamen number, style length and ovule number than most populations of T. teretifolium (Table 2, Figure 2). However, the 7. mengesii plants used to make the crosses had high values for these quantitative traits (Table 2, Figure 2). The 15 sterile F, hybrids were treated with colchicine. Nine of these produced a total of 15 branches with fertile flowers. Flowers from these branches had mean pollen fertility of 94.9% (n= 24), and produced mean fruit set of 50% (n = 40) when used in self and sib crosses. Crosses between fertile flowers of the colchicine-treated F, hybrids and those of Talinum teretifolium plants yielded fertile pro- geny. Ten offspring from these crosses had a mean of 80% pollen fertility and produced an average of 31.2 (s.d., + 7.5) seeds per capsule. Ten open-pollinated T. teretifolium plants had a mean of 85% pollen fertility and produced an average of 32.7 (s.d., + 10.8) seeds per capsule. DISCUSSION The discovery that Talinum parviflorum and T. appalachianum are at the same ploidal level eliminated the major obstacle to con- sidering these one species. The report of 2n = 48 by Steiner (1944) for a single population of 7. parviflorum from Springdale, Arkan- sas, may be a case of autopolyploidy. In addition to having the same chromosome number, 7. appalachianum and T. parviflorum are STAMEN NUMBER 128 Rhodora [Vol. 87 7§ 74 fe) ° os e e e 45 + 4 wt x * OT.m. parent populations wh, @T.m. populations 16 + * T.m./T.p. progeny “ * T.t. populations — AT.p. parent population A AT.p. populations T T qT qT T qT of 2 4 6 STYLE LENGTH (mm) Figure 2. Comparison of synthetic hybrids (open stars) and T. teretifolium (solid stars) for two floral characters which are intermediate between the putative parent species, T. mengesii (circles) and T. parviflorum (triangles). Two T. mengesii popula- tions (open circles) and one T. parviflorum population (open triangle) were used in the interspecific crosses. All symbols denote population means. similar in habit, floral morphology, time of diurnal anthesis and breeding system. Furthermore, fertile hybrids resulted from crosses between the two taxa. These data support the hypothesis that the population which Wolf (1939) named 7. appalachianum should be recognized as 7. parviflorum. Talinum appalachianum is currently listed in the Federal Register (U.S.D.1., 1983) as a proposed endangered species, but one in need of further biological research and field study. The evidence pres- ented in this paper that 7. appalachianum is not a distinct species should assist the Interior Department in deciding if this species is to be included in the permanent Endangered Species list and protected by law. In attempting to reconstruct the evolutionary origin of an allote- traploid, the first consideration was the choice of diploid species to 1985] Carter & Murdy — Talinum 129 be used as the presumed ancestors. Talinum mengesii was chosen as one of the putative parents because a study of meiotic chromosome pairing in triploid and tetraploid hybrids between 7. teretifolium (2n = 48) and 7. mengesii (2n = 24, and 2n = 48) revealed that one-half of the chromosome complement of the tetraploid T. tere- tifolium was homologous with the entire chromosome complement of 7. mengesii (Black & Murdy, 1972). Since interspecific hybrids between Talinum species are intermediate for all major traits that separate species (Murdy, et al., 1970; Krebs, 1971), the only species which could satisfy the extrapolated requirements for the other diploid ancestor was T. parviflorum (T. appalachianum, Alabama) which has small floral structures, few stamens and late anthesis. Talinum calcaricum was eliminated as a possibility because it is a tetraploid species. Ta/inum calycinum was eliminated from consid- eration because its range is far removed from that of T. teretifolium (Figure 1), and it has red flowers (Bogle, 1969) and sepals persistent in fruit (Wilson, 1932). Talinum teretifolium, T. parviflorum and T. mengesii have pink flowers and deciduous sepals. It could be argued that the morphological similarity between the synthetic allopolyploid and Talinum teretifolium has more to do with the choice of parents than it does with genetic relatedness. However, the fact that crosses between the synthetic allotetraploid flowers and T. teretifolium produced fertile offspring is evidence that the genomes of the two entities are at the same ploidal level and are compatible. Fertile hybrids do not necessarily result when crosses are made between Talinum species at the same ploidal level. For example, only sterile hybrids were produced in crosses at the diploid level between 7. mengesii and T. parviflorum. Sterile hybrids also were produced from interspecific crosses at the tetraploid level between 7. teretifolium and T. calcaricum (Krebs, 1971, Master’s thesis, Emory Univ., GA) and between T. reretifolium and an auto- polyploid 7. mengesii (Black & Murdy, 1972). Talinum parviflorum populations in Alabama are separated from their nearest conspecific populations in Arkansas by hundreds of miles essentially devoid of rock outcrops. Do these Alabama popu- lations represent a recent colonization from the west or the last remnant of a once more extensive distribution east of the Missis- sipp! River? Evidence presented in this paper that 7. parviflorum was one of the progenitors of 7. rererifolium, which does not occur west of Georgia, supports the latter. Perhaps the demise of 7. parvi- 130 Rhodora [Vol. 87 florum in the east is related to the spread of its allotetraploid deriva- tive, T. teretifolium. CONCLUSION Several lines of evidence support the conclusion that Talinum appalachianum is not a distinct species, but instead is conspecific with T. parviflorum. Artificial hybridization between 7. mengesii and T. parviflorum produced sterile plants similar in all outward appearances to the alleged allotetraploid, 7. teretifolium. Chromo- some doubling produced fertile tetraploid flowers which, when crossed with T. reretifolium, produced fertile offspring. This result is evidence that the synthetic and natural allopolyploids have compat- ible genomes and supports the hypothesis that 7. reretifolium origi- nated as an allopolyploid derivative of 7. mengesii and T. parviflorum. LITERATURE CITED ANDERSON, E. 1949. Introgressive Hybridization. John Wiley and Sons, New York. Back, C. B., AND W. H. Murpby. 1972. The evolutionary origin of Talinum teretifolium. Bot. Gaz. 133: 405410. BoGir, A. L. 1969. The genera of Portulacaeae and Basellaceae in the southeast- ern United States. J. Arnold Arb. 50: 5665598. MOHLENBROCK, R. H. 1975. Guide to the Vascular Flora of Illinois. Southern Illinois Univ. Press, Carbondale. Murby, W.H. 1968. Plant speciation associated with granite outcrop communi- ties of the southeastern Piedmont. Rhodora 70: 394£407. .T. M. JOHNSON, AND V. K. WRIGHT. 1970. Competitive replacement of Talinum mengesii by T. teretifolium in granite outcrop communities of Georgia, U.S.A. Bot. Gaz. 131: 186&192. STEINER, E. 1944. Cytogenetic studies on Ta/inum and Portulaca. Bot. Gaz. 105: 3745379. U.S. DEPARTMENT OF THE INTERIOR. 1983. Endangered and threatened wildlife and plants; supplement to review of plant taxa for listing as endangered or threatened species. Federal Register 48: 53640£53670. Ware, S. 1967. A new Talinum (Portulacaceae) from the cedar glades of middle Tennessee. Rhodora 69: 4665475. Witson, P. 1932. Talinum. North American Flora 21: 280£289. Wor, W. 1939. The status of Ta/inum in Alabama. Amer. Midland Nat. 22: 315§332. DEPARTMENT OF BIOLOGY EMORY UNIVERSITY ATLANTA, GA 30322 PANAX QUINQUEFOLIUM (ARALIACEAE) IN SOUTHEASTERN OKLAHOMA WALTER H. LEWIS AND GARY A. THOMPSON In the western ridges of the Ouachita Mountains a number of interesting range extensions of more northeastern vegetation are occasionally found. An example of such a find is Panax quinquefo- lium L. (American ginseng), a species rare in hilly, deciduous woods of two Oklahoma counties, Leflore and McCurtain. The following collections represent the most southwestern distribution of the species. OKLAHOMA: Leflore Co.: 5 miles S of Big Cedar, 7 Oct 1973, Bradley 283 (DUR, NLU, VDB); Page, 9 Sep 1913, Stevens 2757 (Ny); Rich Mt., ca. 7 miles from ject. USI and US259, 25 Sep 1979, Taylor 28244 (DUR, VDB); Rich Mt., 8.5 miles W of Arkansas-Oklahoma state line on Hwy. |, 2 Aug 1984, Lewis & Thompson 9805 (DUR, MO, OKL). McCurtain Co.: 8.4 miles NE of Smithsville, 11 Oct 1981, Magrath 12349 (OCLA). We Hi; &. DEPARTMENT OF BIOLOGY WASHINGTON UNIVERSITY ST. LOUIS, MO 63130 G. A. T. DEPARTMENT OF HORTICULTURE UNIVERSITY OF WISCONSIN MADISON, WI 53706 131 132 Rhodora (Vol. 87 IN MEMORIAM Dr. James P. Poole, a member of the New England Botanical Club since 1945, died December I 1, 1984, in Hanover, New Hamp- shire at the age of 95. A professor emeritus of botany at Dartmouth College, he had taught there for 35 years. He continued to serve after retirement in 1957 as the curator of the Jesup Herbarium. Born in Gloucester, Mass., he earned his BS in forestry from the University of Maine in 1912, his master’s degree in 1918 and his doctorate in Botany in 1921 from Harvard University. Dr. Poole was recently voted by the Council to be an Honorary Member of the Club. He had written to the Club indicating that he could no longer read Rhodora, and because of that fact, felt he should be deleted from our rolls. The Council’s action naming him an Honor- ary Member, to receive Rhodora for life, occurred less than a month before his death. REDISCOVERY OF MALAXIS PALUDOSA (L.) SW. (ORCHIDACEAE) IN MINNESOTA TIMOTHY REEVES AND LINDA REEVES ABSTRACT The first documented field observation of Ma/laxis paludosa in Minnesota since 1938 is reported. Herbarium specimens confirm the occurrence of the species in five counties in the state; three represent newly published records. The species occurs in Alaska, Canada and Eurasia. Description of the habitat and list of associated species is provided. One site is reported where M. pa/udosa grows sympatrically with its congeners M. unifolia and M. monophyllos. Key Words: Orchidaceae, Malaxis paludosa, bogs, Minnesota Malaxis paludosa (L.) Sw. is a circumboreal orchid known in the continental United States only in Minnesota (Correll, 1950; Fer- nald, 1950; Gleason & Cronquist, 1963; Case, 1964; Luer, 1975; Petrie, 1981; Marquis & Voss, 1981). A survey of the literature and herbarium specimens (F, MICH, MIL, MIN, MSC, SDC, UWM, WIN, and WIS) revealed no Minnesota collections since 1938. The fact that two recent comprehensive photographic guides to North American Orchids (Luer, 1975; Petrie, 1981) have photographs of the species taken in Alaska and/or Canada, suggests that the Min- nesota populations are infrequently seen. Our examination of spec- imens and field explorations since 1981 have added to the known distribution of the species in Minnesota. Other recently located sites found by Minnesota Department of Natural Resources personnel are also reported. This inconspicuous orchid is known from scattered localities in Eurasia and North America. In Canada the species is known from British Columbia, Alberta, and Ontario. An excellent account of the Canadian localities is given by Baldwin (1961). We are aware of previously unreported collections from Manitoba (N of Gilbert Plains, 9 Aug 1970 and 18 Aug 1971, Parker s.n., WIN, G. M. Keleher, pers. comm.). The United States collections are from Alaska and Minnesota. The Alaska localities are listed by Baldwin (1961). Previous collec- tions in Minnesota have come from the following counties: Otter Tail Co., Jul 1904, Lyon s.n. (MIN!); Clearwater Co., Brayton 3165 133 134 Rhodora [Vol. 87 (MIN!), Grant 3155 (MIN!, NY!), Moyle 1266 (MIN!), Rosendahl 352 (US!), Rosendahl! 4837, 5930, 5941 (MIN!); Hubbard Co., 11 Jul 1934, Moyle s.n. (MIN!), 29 Aug 1934, Rosendahl 6957 (MIN!) [These Hubbard Co. locations are labelled “Clearwater Co.” but cited localities are in Hubbard Co. Paul Rundell (pers. comm.) reported that the species was seen in the area in the 1970’s.]; Becker Co., 11 Jul 1934, Moyle 1450 (F!). This report is apparently the first published one of the Hubbard and Becker Co. collections. The spe- cies was reported from Itasca County by Correll (1950) but we cannot locate substantiating specimens. On July 26, 1981, we located a population of Malaxis paludosa in Beltrami County, Minnesota (7. & L. Reeves 7336 with A. Schmierer, MIN, NY, GH). This is the fifth county in which the species has been documented in the state. This site (which will not be identified precisely to protect the plants) is in a boggy conifer swamp. The dominant trees are Picea mariana (Mill.) BSP., Abies balsamea (L.) Mill., and Larix laricina (DuRoi) K. Koch. Common shrubs are A/nus rugosa (DuRoi) Spreng. and Ledum groenlandi- cum Oeder. Betula pumila L. occurs in the area. Malaxis paludosa usually grows in semi-shaded sites which receive some sunlight during part of the day. The orchid flowers from late June to early August; fruiting plants persist into Sep- tember. The orchid typically occurs on a mossy hummock, 15-30 cm above the water level in adjacent wet depressions. Plants have been found at the base of black spruce, balsam fir, and alder as well as on hummocks not near a trunk. Sphagnum carpets the forest floor but the plants of M. paludosa are most frequent on other mosses (but sometimes grow on Sphagnum). Commonly associated mosses are Pleurozium schreberi (Brid.) Mitt., Thuidium delicatu- lum (Hedw.) B.S.G., Helodium blandowii (Web. & Mohr) Warnst., Callicladium haldanianum (Grev.) Crum, Pohlia nutans (Hedw.) Lindb., Plagiomnium ciliare (C. Mull.) Kop., and Sphagnum sp. Commonly associated herbaceous species are: Carex disperma Dewey, Rubus pubescens Raf., Viola canadensis L., Mitella nuda L., Fragaria vesca L., Pyrola secunda L., Linnaea borealis L., and Moneses uniflora (L.) A. Gray. Other herbaceous species found near M. paludosa are Malaxis unifolia Michx., Coptis trifolia (L.) Salisb., Aralia nudicaulis L., Platanthera obtusata (Banks ex Pursh) Lindl., Listera cordata (L.) R. Br., Goodyera repens (L.) R. Br., 1985] Reeves & Reeves — Malaxis 135 Circaea alpina L., Epilobium palustre L, Gaultheria hispidula (L.) Muhl., Trientalis borealis Raf., Galium aparine L., Galium obtu- sum Bigel, Thelypteris palustris L, Equisetum fluviatile L., Botry- chium virginianum (L.) Sw., and Dryopteris cristata (L.) A. Gray. The general area also has Malaxis monophyllos (L.) Sw., Ameror- chis rotundifolia (Banks) Hulten, Cypripedium reginae Walt., Cypripedium acaule Ait., Platanthera hyperborea (L.) Lindl., Pogonia ophioglossoides (L.) Ker., Calopogon tuberosus (L.) BSP., Corallorhiza trifida Chat., Sarracenia purpurea L., Drosera rotun- difolia L., Caltha palustris L., Menyanthes trifoliata L., Ribes lacus- tre (Pers.) Poir., Eriophorum viridi-carinatum (Engelm.) Fern., and Aralia nudicaulis L. In addition to the Beltrami County locality discussed above we are aware of undocumented occurrences in Lake of the Woods (two sites) and Cass Counties, two other localities in Hubbard Co., one other site in Becker Co., and two additional Beltrami Co. localities (Paul Rundell, pers. comm.). We have located only a few plants in three years of searching. In 1981 we located [5 plants in several hours of searching. In 1983 we found only 37 plants although we spent many hours throughout July looking for plants. Our field work in 1984 has increased the total number of plants seen to nearly 50. The total area searched at this site is about 65 ha and only a few scattered plants have been seen any distance from the main population. The diminutive plants of Malaxis paludosa commonly possess foliar embryos on leaf apices similar to those described and discussed by Taylor (1967). At one site three species (M. paludosa, M. unifolia, M. mono- phyllos) occur together. Our observations indicate that the flower- ing periods of the three species overlap considerably. This site is the only one of sympatry that we have seen and we are not aware of any other such site in North America. We are currently in the second year of a study of the pollination biology of M. paludosa, M. unifo- lia, and M. monophyllos. A report of our research on M. paludosa has been published in the American Orchid Society Bulletin (Reeves & Reeves, 1984). ACKNOWLEDGMENTS We thank William R. Buck for the moss determinations, Alan Schmierer for valuable field assistance and the entire Schmierer 136 Rhodora [Vol. 87 family for extensive support. We also thank Paul Rundell of the Minnesota Department of Natural Resources for his considerable support, information and advice, Russell Simmons and John Fyl- paa for their interest, and the curators of the above-cited herbaria for their courtesy. LITERATURE CITED BALDWIN, W. K. W. 1961 Malaxis paludosa (L.) Sw. in the Hudson Bay low- lands. Canad. Field—Naturalist 75; 74-77. Cask, FRED W., JR. 1964 Orchids of the Western Great Lakes Region. Cranbrook Inst. Sci., Bloomfield Hills, MI. CoRRELL, DONOVAN S. 1950. Native Orchids of North America North of Mexico. Chronica Botanica Company, Waltham, MA. FERNALD, M. L. 1950 Gray’s Manual of Botany. 8th ed. American Book Com- pany, Boston. GLEASON, HENRY A. and ARTHUR CRONQUIST. 1963 Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Van Nostrand Company, N.Y. LueR, CARLYLE A. 1975. The Native Orchids of the United States and Canada Excluding Florida. N.Y. Bot. Garden, N.Y. Marquis, RoBerT J. and Epwarp G. Voss. 1981. Distributions of some western North American plants disjunct in the Great Lakes region. Michigan Bot. 20: 53-82. PETRIE, WILLIAM. 1981. Guide to Orchids of North America. Hancock House, Blain, WA. Reeves, LINDA M., and Timotuy Reeves. 1984. Life history and reproduction of Malaxis paludosa in Minnesota. Amer. Orchid Soc. Bull. 53(12): (in press). Taytor, Roy L. 1967. The foliar embryos of Malaxis paludosa. Canad. J. Bot. 45: 1553-1556. DIVISION OF SCIENCE AND MATHEMATICS UNIVERSITY OF MINNESOTA, MORRIS MORRIS, MN 56267 STUDIES IN THE ARISTIDA (GRAMINEAE) OF THE SOUTHEASTERN UNITED STATES. I]. MORPHOMETRIC ANALYSIS OF A. INTERMEDIA AND A. LONGESPICA' KELLY W. ALLRED ABSTRACT Morphologic features of the spikelets of Aristida intermedia Scribn. & Ball and A. longespica Poiret were compared using principal components and stepwise discrimi- nant analyses. Major phenetic overlap existed between the two taxa. Central awn and lateral awn lengths were the best discriminators angle could not be used effectively. A single species with two varieties is proposed, var. /ongespica and var. geniculata (Raf.) Fern. Key Words: Aristida intermedia, A. longespica, principal components, taxonomy, stepwise discriminant analysis In an attempt to provide the treatment of Aristida for the Vascu- lar Flora of the Southeastern United States project, several prelimi- nary taxonomic surveys of this difficult genus were undertaken. The first (Allred, 1984) examined the morphologic relationships of A. purpurescens Poiret, A. tenuispica Hitchcock, and A. virgata Tri- nius. In this study, phenetic affinities of A. intermedia Scribner & Ball and A. /ongespica Poiret are examined. These surveys provide a justification for the taxonomic judgments that will be rendered in the floristic treatment, and are preliminary to more complete biosys- tematic investigations. Aristida longespica was named from material collected in the Carolinas by Bosc. It is rather common throughout the eastern United States from New Hampshire to Florida and westward to Texas, Kansas, and lowa. Aristida intermedia was named from a Mississippi collection made by Kearney, and is more common in the western range of A. /ongespica. Scribner and Ball drew the name intermedia from their belief that it was nearly intermediate between A. longespica and A. purpurescens. Both A. longespica and A. intermedia are erect annuals with often extensive vegetative branch- ing and terminal, sometimes full, panicles. The two are usually 'Journal Article No. 1084 of the New Mexico Agricultural Experiment Station, Las Cruces, New Mexico. 137 138 Rhodora [Vol. 87 separated using dimensions of the glumes, lemmas, and awns, with A. intermedia being larger. Gould (1975) regarded the two as only varietally distinct and applied the varietal name geniculata (Raf.) Fernald to the A. intermedia taxon. The purposes of this study were to examine the morphologic vari- ability of Aristida intermedia and A. longespica, particularly in the southeastern United States, and to determine the proper taxonomic status for any meaningful morphologic entities perceived. MATERIALS AND METHODS After careful study of the original descriptions and descriptions of the types in Henrard (1927), 172 specimens from throughout the presumed range of both taxa were selected for analysis, with about half thought to correspond to each taxon. However, no firm a priori identifications were made, and intermediate or problematic speci- mens were included. Three specimens of Aristida dichotoma Michaux were also included for reference to a closely related spe- cies. To develop OTUs (operational taxonomic units), each speci- men was scored for the following characteristics: length of glumes, lemma, callus, and awns; and angle of deflection of the awns. These features are used to distinguish A. intermedia and A. longespica, but not necessarily A. dichotoma. Vegetative features were judged to be so similar as to be non-discriminatory and were not measured. A single mature spikelet from the upper third of the panicle was measured. The data from each OTU were standardized so that each variable had a standard deviation of 1.0; they were subjected to principal components and stepwise discriminant analyses using the BMDP statistical package (Dixon, 1981). RESULTS AND DISCUSSION The principal components analysis (PCA) was based on a varia- ble-by-variable correlation matrix. Variables with the highest correlations were central awn length and lateral awn length (r = 0.868), glume I length and glume II length (r = 0.762), and central awn length and lemma length (r = 0.744). These variables ought to be the most effective discriminators for these OTUs, if groups are to 1985] Allred — Aristida 11 139 be discriminated. Central and lateral awn angle, considered diag- nostic by many authors, had low correlation values; none was greater than + 0.280. The first and second components accounted for 55% and 14%, respectively, of the variation present in the study sample. Dispersion of OTUs along Component | was a function of size and reflected correlations with the length variables. Component II accounted for variability associated with awn angles (Table I). The projection of OTUs onto the PCA grid showed no clear-cut separation into phenetic groups, with the exception of the three dichotoma OTUs located in the extreme “northwest” quadrant (Figure 1). Although spikelets of dichotoma and longespica appeared to be very similar (disregarding the coiled awn of dicho- toma, which was not evaluated in this study), the PCA suggested that dichotoma was phenetically distinct from the /ongespica and intermedia entities. Examination of specimens revealed that OTUs with longer awns, longer lemmas, and longer glumes (intermedia) tended to be located in the “eastern” portion of the PCA grid, and that OTUs with shorter awns, shorter lemmas, and shorter glumes (/ongespica) were placed in the “western” portion. Although the PCA demonstrated that size features of the spikelet, especially awn lengths, contributed to the dispersion of the OTUs along a size gradient of long to short, it is not an identification procedure and did not identify which OTUs belonged to intermedia and which to /ongespica, nor where the phenetic boundary between the two lay. Table I. Factor loadings for components I and Il in the PCA of Aristida longespica and A. intermedia OTUs. Loadings less than 0.250 are replaced by zero. Variable Component | Component II Central awn length 0.887 0.0 Lateral awn length 0.882 0.0 Glume | length 0.869 0.0 Glume II length 0.852 0.0 Lemma length 0.832 0.0 Callus length 0.763 0.0 Lateral awn angle 0.0 0.714 Central awn angle 0.0 0.597 140 Rhodora [Vol. 87 3 +, oe O 2 r i) 7 o fe] 5 o 90 t) ° Ww oO oo 2 - fone) eg 1 0° 0 OPo00 cy = oO fe) re) re) 090 900 tome) 8 Pf 520% 00 90 P0 00 0° fe} o 090 So oo 0% ° J+ 9 7 9 9 90000° 0 00 °