NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 14 (3) 2008

IN THIS ISSUE:

RELOCATING TERRESTRIAL ORCHID PLANTS ALIEN TERRESTRIAL ORCHID, EULOPHIA GRAMINEA, INVADES MIAMI GENERAL NOTES ON THE ECOLOGY OF SACOILA PALUDICOLA, SARASOTA COUNTY, FLORIDA THE TAXONOMY AND DISTRIBUTION OF SPIRANTHES CASEI IN NORTHERN NEW ENGLAND

and more………….

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

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NORTH AMERICAN NATIVE ORCHID JOURNAL
Volume 14 (3) 2008

CONTENTS NOTES FROM THE EDITOR 185 RELOCATING TERRESTRIAL ORCHID PLANTS Robert J. Ferry, Ph.D. 187 ALIEN TERRESTRIAL ORCHID, EULOPHIA GRAMINEA, INVADES MIAMI Bob Pemberton, Ph.D., Suzanne Koptur, Ph.D. & Timothy Collins, Ph.D. 190 THE GENUS SACOILA (ORCHIDACEAE) IN THE CONTINENTAL UNITED STATES AND A NEW COMBINATION Paul Martin Brown 192 GENERAL NOTES ON THE ECOLOGY OF SACOILA PALUDICOLA, SARASOTA COUNTY, FLORIDA Craig N. Huegel, Ph.D. & Kathleen K. McConnell 200 THE TAXONOMY AND DISTRIBUTION OF SPIRANTHES CASEI IN NORTHERN NEW ENGLAND with references to extant sites in New York, Pennsylvania, and Nova Scotia Paul Martin Brown 212 WHATEVER GOT YOU STARTED IN ORCHIDS? The Slow Empiricist 234
Unless otherwise credited, all drawings in this issue are by Stan Folsom. The opinions expressed in the Journal are those of the authors. Scientific articles may be subject to peer review and popular articles will be examined for both accuracy and scientific content. Volume 14 (3): 185-236 issued July 15, 2008. Copyright 2008 by the North American Native Orchid Journal Cover: Epidendrum floridense by Stan Folsom

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NOTES FROM THE EDITOR It seems all too often that well-laid plans get sidetracked and such is the case with this issue. Several of the articles planned for this issue will appear in October, primarily because they will contain more information that will reflect the 2008 field season, as well as because some of them were just not prepared in time for this issue. But, the good news is we have a wide variety of excellent articles from several authors. My paper on Spiranthes casei in Northern New England is presented as a request item from several Spiranthes enthusiasts. It was the basis of my Master‘s thesis in 1997 and I have included notes to bring some of the information up to date. The electronic format continues to be well received and we now reach more than 1200 readers. You may read back issues at: http://wiki.terrorchid.org/tow:journals. The current update of the North American Personal Checklist is also available at that website. The checklist will be updated quarterly as needed. Paul Martin Brown Editor naorchid@aol.com 10896 SW 90th Terrace, Ocala, FL 34481 PO Box 759, Acton, Maine 04001 June-mid October Coming in October….. A NEW SPECIES OF FRINGED PLATANTHERA FROM EASTERN NORTH AMERICA ADDITIONAL DISTRIBUTIONAL INFORMATION REGARDING SPIRANTHES STELLATA WILD ORCHIDS IN THE 21ST CENTURY: RECENT PUBLICATIONS NEW COLOR FORMS FROM THE CENTRAL APPALACHIANS AND MUCH MORE….

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RELOCATING TERRESTRIAL ORCHID PLANTS
Robert J. Ferry, Ph.D.
Reprinted from McAllen International Orchid Society Journal 9(5): 6-9. 2008. Introduction: In general, the advice regarding terrestrial orchids are to leave them where they are. Don‘t pick the flowers! Unless you‘re well-equipped to propagate them from seed, don‘t collect seed! Above all, try to preserve the habitat! The advice is observe them (watch where you step!), photograph, and try to spot and keep a record of any pollinators, but-in short-leave the plants undisturbed. However, given the march of human proliferation and the building of housing developments (which is usually euphemistically termed ―progress‖), what is the conservationist and scientist to do when a habitat is slated to have ―progress‖ enter and convert a habitat into a development, an athletic field, or for some other such use by an ever-increasing number of humans? One answer is to legally stop the so-called ―progress.‖ Then again, the danger to the plant or animal species may be from local governmental agencies unable (or unwilling) to prevent local individuals from illegally harvesting (killing) the species. What can be done to preserve, conserve, propagate, and proliferate the species? This piece doesn‘t address the conservation avenues for animals (e.g. tigers, elephants, and the like). However, this piece does offer one approach to relocating relatively small, herbaceous plants. No, it‘s not presented as an avenue for the hobbyist-fanatic to ―dig-and-haul-away!‖ The words of this essay‘s first paragraph still apply, but-faced with the reality of a population‘s extermination-the following protocol may be useful in relocating endangered plants whether by colleagues in the United States, Canada, Mexico, India, South Africa, or elsewhere worldwide. Background: Previous issues of this publication have dwelt on Spiranthes parksii, an endangered United States terrestrial orchid. In the College Station area of Texas, a forested habitat of this orchid is slated to be converted to use as a landfill, but the destruction has been delayed until plants of S. parksii can be relocated. When your editor visited this area, Loblolly Pine (Pinus taeda L.; Oldfield Pine), Red Cedar (Juniperus virginiana L.; Cedar, Savin, Red Juniper), and other trees and shrubs were noted. Plants of Spiranthes cernua, a terrestrial orchid not on the endangered list was also observed. Listing all the area‘s species would be a somewhat lengthy flora in itself, but as far as is known, the only species slated for recovery and relocation is the endangered S. parksii. To successfully relocate this terrestrial orchid, it is necessary to obtain, relatively undisturbed, the individual plant and its root-network of tubers. As is very well known among orchidists interested in terrestrials, bare-root transplanting is a near-certain way to kill the plant! A way must be devised to obtain the plant and its immediate soil-fungi system without
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disturbing plants, and care must be taken to relocate them into ecologically friendly habitats! Ryan Hammons, a Masters student under Dr. Fred Smiens of Texas A & M University is doing just that, and with his cooperation and that of Dr. Smeins, your editor was able to witness and photograph part of the operation as a part of efforts of the committee assembled to oversee the recovery, relocation, and proliferation of this species. Many individuals have been very cooperative in the common thrust on behalf of Spiranthes parksii, and some excellent field work continues to be done by graduate students Linda Langlitz and Ryan Hammons. It continues to be a pleasure for this individual to work with these individuals and other members of the Recovery team. Materials and Methods: One needs a sturdy plastic ring about 30 cm in diameter and about 20 cm in height. This may be cut from a section of water or sewer line. The main requirement for the plastic section is that it be sturdy (rigid) enough to withstand some serious hammering without cracking or shattering. Also needed will be a stout hammer for hammering the ring into the soil. Once the ring is embedded in the soil surrounding the plant, a good spade or a flat shovel comes into play. After an entry notch is cut alongside one edge of the ring, the flat shovel is slid beneath the ring and the plug of soil, with the plant, is lifted out of the ground. A suitable pot is now needed, and the commercial 12-inch (22-23 cm) ―bulb pan‖ plastic pot fills the need admirably. The plugand-plant are eased into the pot and voila! The plant is ready to be transported. One should also have either a set of plant tags or some type of permanent marking pen to mark individual pots as plant-plugs are taken. In addition, a pencil-and-paper field record should be kept of the plant showing from where it was taken and what disposition is being made of it. With regard to this last item, a detailed computer record is strongly indicated. Discussion: Note the plastic ring and hammer alongside the worker (Fig. 1). In the figure, the blue flags represent plants identified and confirmed as S. parksii. Orange flags and white flags indicate other plants, some of which are S. cernua. The time for such recovery work is obviously during relatively dry weather as in rainy weather (or shortly thereafter) the ground would present a much more difficult working environment as well as make the taking of a plug a sloppy and generally messy task!

Fig. 1. Ryan Hammons in the field, with equipment.
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Looking more closely at the ―nuts-andbolts‖ of the operation, we see the plastic ring hammered into the ground (Fig. 2, top left). With the ring in the ground, a key-hole plug is removed (Fig. 3, center left). With the access hole available, the flat shovel is inserted beneath the plastic ring holding the plant and soil plug, and the plug is then gently deposited in a pot for transport (Fig. 4, bottom left). As shown, it‘s all a very simple task. However what is not necessarily shown is the traversing of land that is sometimes a bit rougher than one‘s front lawn! In addition, if several soil plugs are taken and potted, returning them to the faithful Jeep (seen in Fig. 5), can be something more than a simple walk in the park! For example, note the gulley on the way back to the Jeep (Fig. 6). Incidentally, the faithful Jeep which has figured in so many expeditions in the United States, Canada, and Mexico, had turned over just a little more than 300,300 miles as it‘s seen in Figs. 5 and 6! It brought Ryan and the author out of the ‗boondocks‘, back to the university, and then transported your editor back to another part of town where he met with Linda Langlitz prior to his intended departure back to the Victoria area. Strange noises then came from the Jeep‘s engine! Ultimately Linda Langlitz kindly transported your editor back home, some 160 miles distant, and we arrived home at about 2200 hours! Mechanic Bubba and your editor returned to College Station a few days later with a truck and trailer, and hauled the Jeep back to the Victoria area for repairs. The word is that a nearly-new engine is being ordered, and then, hopefully, the Jeep will be good for at least another hundred to hundred and fifty thousand miles. At least the Jeep lasted long enough to get us out of the
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boondocks! Our situation that day could have been much worse! In closing, make no mistake: this piece is not on how to steal terrestrial orchid plants! If the habitat can be conserved, that‘s the place for the plants; not in some personal collection! Serious orchidists and conservationists take note!

Fig. 5. The ring is lifted from the ground

Fig.6. Typical deep erosion-caused gully. Robert J. Ferry, Ph.D., 343 John Wayne Trail, Victoria, Texas 77905 rferry@miosjournal.org

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ALIEN TERRESTRIAL ORCHID, EULOPHIA GRAMINEA, INVADES MIAMI
Bob Pemberton, PhD, Suzanne Koptur, PhD & Timothy Collins, PhD

We first encountered an Asian orchid, Eulophia graminea, in South Miami during the autumn of 2007. Thus far we have found it in seven residential areas and in two parking lot islands, one at a supermarket in northern Miami and the other adjacent to the beach on Virginia Key in Biscayne Bay in Miami-Dade County in southeastern Florida. The area where we have detected plants of this orchid stretches 35 km from north to south. The orchid appears to have been at some locations for two years or more. Eulophia graminea is native to the tropical and subtropical parts of Asia from Pakistan east through India, Nepal, Southeast Asia, southern China to the Ryuku Islands south of Japan. It also extends into cooler areas in Kashmir in northwestern India and Sikkim a Himalayan kingdom in northeastern India. In eight of the nine sites where we have found the orchid, it is growing in woodchip mulch. In its native range the plants grow in many kinds of open, disturbed habitats, including grasslands and even beaches. The small flowers (Figure 1), usually about 2.5 cm across, are not showy but are attractive when viewed close up, with a white lip is marked with rose-pink, contrasting nicely with the somber green petals and sepals. The inflorescences (Figure 2) arise from spherical to conical pseudobulbs, usually about 5-8 cm in diameter, which typically sit completely or partly above the ground. The slender inflorescences range from 30 cm to 1.5 m tall and bear up to 60 flowers. In addition to defining the naturalization, we have been studying the orchid‘s reproduction. Most of the plants are producing fruit but at low levels, and the flowers appear to need a pollinator. The flowers are faintly fragrant and bear a small nectar-filled spur, attractants probably for an insect pollinator. Determining what the pollinator(s) and the mycorrhizal fungi that are promoting the germination of its seed and seedling establishment are the focus of our research. We suspect that Eulophia graminea entered Florida or the United States as plants imported for cultivation. Plants are offered for sale by nurseries in Thailand via the internet. Amateur growers outside the United States also offer to trade the orchid via the internet. The orchid then probably escaped cultivation to naturalize, a many horticultural plants but relatively few orchids have done. The naturalized orchid plants can spread via its airborne seed and pseudobulbs moved in mulch and soil. The northern parts of its native distribution in Kashmir and Sikkim have colder climates than southern Florida‘s subtropical climate, which suggest that the orchid will be able to survive farther north in Florida and perhaps beyond. The orchid‘s airborne seed may spread it south into the West Indies. People may purposefully move the orchid for cultivation Eulophia graminea almost certainly occurs in many more places than where we have detected it. We wish to more accurately define its current distribution and to monitor its spread. We request your help in this process. If you encounter plants, please send us photos or
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specimens and some details of the location and occurrence. For additional information see our recent technical paper ―An Asian orchid, Eulophia graminea (Orchidaceace: Cymbidiae), naturalizes in Florida‖ Lankesteriana (International Journal of Orchidology) (2008) vol. 8: 5-14, and our popular article of the same topic ―Newly naturalized orchid (Eulophia graminea) found in Florida‖. Orchids (2008) 77: 412-13.
Bob Pemberton, PhD, Fairchild Tropical Botanic Garden, 2121 SW 28th Terrace, Ft. Lauderdale, FL 33312, USA. (e-mail Robert. Pemberton@ars.usda.gov). Suzanne Koptur, PhD and Timothy Collins, PhD, Department of Biological Sciences, Florida International University, Miami, FL 33199 USA (e-mails kopturs@fiu.edu; Collinst@fiu.edu).

Fig. 2 left

Fig. 1 below

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Fig. 1 Sacoila lanceolata (Aublet) Garay

Fig. 2 forma albidaviridis Catling & Sheviak Fig. 3 forma folsomii P.M. Brown

Fig. 4 Sacoila paludicola (Luer) P.M. Brown

Fig. 5 forma aurea P.M. Brown

Fig. 6 Sacoila squamulosa (Kunth)Garay

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THE GENUS SACOILA (ORCHIDACEAE) IN THE CONTINENTAL UNITED STATES AND A NEW COMBINATION
Paul Martin Brown The genus Sacoila in the continental United States, restricted to Florida, for many years was considered to be a single, somewhat variable, species within the genus Spiranthes (revalidated in the genus Sacoila by Garay (1982). Carlyle Luer, in 1971, described a new variety of that species as var. paludicola, a rather unique taxon from the swamps of southwestern Florida. In 2000, plants of Sacoila squamulosa, a species known from the West Indies, were discovered in Marion County in central Florida. In addition several interesting color forms have also been described from the S. lanceolata complex. The recent discovery of a large indigenous colony of S. lanceolata var. paludicola in Sarasota County, Florida has led to renewed interest in the genus and specifically the status of that taxon. With more evidence now available it is here proposed at the species level as Sacoila paludicola (Luer) P.M. Brown. Key to the taxa of Sacoila in the United Florida 1a flowers/ovaries with prominent scurfy or hoary dots; mentum ca. 2/3 as long as the ovary…S. squamulosa 1b flowers/ovaries lacking hoary dots; mentum ca. ½ as long as the ovary…2 2a leaves always absent at flowering time (late April-mid June); flowers brick-red to terracotta (rarely white/green or bronzy-yellow) apices of petals and sepals divergent to recurved; plants of open roadsides, fields, pine flatwoods; plants apomictic, flowers longlived (up to 2 weeks) …S. lanceolata leaves usually present (to some degree) at flowering time (late Jan.-early April); flowers bright red (rarely yellow) apices of petals and sepals barely divergent; plants of hydric hammocks restricted to southern Florida and western Cuba; flowers autogamous, shortlived (3-4 days)…S. paludicola

2b

Descriptions are based, in part, upon those by Brown and Catling in FNA, 2002. Sacoila lanceolata (Aublet) Garay, Botanical Museum Leaflet 28: 352. 1982. (Figs. 1, 8) Basionym: Limodorum lanceolatum Aublet. Hist. Pl. Guiane 2: 821. 1775 Synonyms: Gyrostachys aphylla (Hook.) Kuntze 1891; Gyrostachys lanceolata (Aubl.) Kuntze 1891; Gyrostachys orchioides (Sw.) Kuntze 1891; Gyrostachys stenorrhynchus Kuntze 1891; Ibidium cristalligerum Salisb. 1812;; Neottia aphylla Hook. 1828; Neottia lanceolata (Aubl.) Willd 1805; Neottia orchioides (Sw.) Willd. 1799; Neottia plantaginea Hkr. 1827; Sacoila lurida Raf. 1836; Satyrium orchioides Sw. 1788; Serapias coccinea Vellozo 1827; Spiranthes jaliscana S. Watson 1891; Spiranthes lanceolata (Aubl.) León 1946; Spiranthes orchioides (Sw.) A. Rich. 1850; Spiranthes orchioides (Sw.) Hemsl. 1884; Stenorrhynchos aphyllus (Hook.) Lindl 1840; Stenorrhynchos australis Lindl. 1840; Stenorrhynchos jaliscana (S. Watson) Nash 1895; Stenorrhynchos orchioides (Sw.) Richard ex Sprengel 1826; Stenorrhynchos orchioides (Sw.) Richard ex Sprengel var australis (Lindley) Cogn. 1895; Stenorrhynchos orchioides (Sw.) Richard ex Sprengel var plataginea (Hkr.) Lindley 1840 Several of the Central and South American taxa included in the synonyms need to be carefully examined to correctly determine their taxonomic status.
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Plants 23–60 cm. Stems: sheaths 5–9, cauline, bladeless. Leaves 2–7, usually absent at flowering time; petiole not distinct; blade elliptic, 5–25(–35) × 2–8 cm. floral bracts green to reddish brown, lanceolate, 15–20(–25) mm. Inflorescences 43–18 cm; rachis pubescent with bubbleshaped, glandular-capitate, and transitional hairs; floral bracts green to reddish-brown, lanceolate. Flowers usually coral, brick, terra-cotta or orange-red, occasionally pale green or golden bronze; dorsal sepal dorsal sepal free 19–23 × 7–9 mm, ascending at apex;; lateral sepals 23–31 × 3–8 mm, connate for 5–7 mm over basal portion of mentum, free portion of mentum 4–6 mm; petals 21–26 × 6–8 mm; lip pale red or pinkish white to white, 23–31 × 8–11 mm; column 9–10 mm from attachment of dorsal sepal to tip, foot extending back 11–12 mm; pollinaria 8–9 mm; viscidia 4–5 mm; pedicellate ovary 10–20 mm; rostellum 4–5 mm. Capsules ca. 14 × 8 mm. ascending, ovoid, expanded portion 9–16 × 6–10 mm. Seeds 0.5–1.4 mm largely polyembryonic. 2n = 46. Fla.; Mexico; West Indies; Central America; northern South America. Sacoila lanceolata is widespread in tropical and subtropical America. In Florida it is found in most counties in peninsular Florida with a disjunct record for Walton Co. in the central panhandle. Synonyms for Sacoila lanceolata in the restricted sense are given by Garay (1980[1982]) and in the very broad sense by Luer (1972). Two color forms associated with S. lanceolata have been named: forma albidaviridis Catling & Sheviak (Fig. 2), with green and white flowers, and forma folsomii P.M. Brown (Fig. 3) with golden bronze flowers. In Florida Sacoila lanceolata is apomictic and produces seed by adventitious embryony. Plants of Sacoila lanceolata from Florida referred to Spiranthes lanceolata var. luteoalba (Reichenbach) Luer (1972) represent only a very restricted clone with more ascending flowers whose characters intergrade with those of plants from other parts of the range. Because all green-flowered plants addressed by Luer do not consistently possess the distinctive features he noted and do not conform to the description of var. luteoalba (Reichenbach f.) Luer, they are best recognized at the rank of forma as forma albidaviridis Catling & Sheviak (1993).

Sacoila paludicola (Luer) P.M. Brown comb. & stat. nov. (Figs, 4, 7, 10)

Basionym Spiranthes lanceolata (Aublet) León var. paludicola Luer, Florida Orchidist 14: 19. 1971. Synonyms: Sacoila lanceolata var. paludicola (Luer) Sauleda, Wunderlin & B. F. Hansen Stenorrhynchos lanceolatum (Willd.) Rich. var. paludicola (Luer) W.J. Schrenk Stenorrhynchos cinnabarinum Lindl. var. paludicola (Luer) W.J. Schrenk Swamp beaked orchid; Fakahatchee beaked orchid Plants 20-40 cm tall; leaves usually present at flowering time. Inflorescences 3–15 cm; floral bracts red, lanceolate, 12–15 mm. Flowers 9–29, deep red (rarely yellow); dorsal sepal 9–11 × 3.5–4.5 mm; lateral sepals 12–14 × 33–42 mm, connate 1–2 mm over basal portion of mentum, distinct portion of mentum 0.8–1.2 mm; petals 9–10.5 × 2.2–2.8 mm; lip red to pale red, 11–13 × 3.8–4.5 mm; column 6–7.5 mm from attachment of dorsal sepal to tip, foot extending back 3– 4 mm; pollinaria 3–3.5 mm; viscidium 2.9–3.1 mm; pedicellate ovary 12–15 mm; rostellum 2.3– 2.8 mm. Capsules 12 × 6 mm. Seeds monoembryonic. Flowering late Jan-early Apr. Moist to wet hardwood forests and palmetto hammocks on sandy or organic substrates; Florida; western Cuba Sacoila paludicola produces seed by auto-pollination (Catling, 1987) and is very uncommon and with native colonies known only from Collier and Sarasota counties, Florida. Plants have been introduced in Miami-Dade, Palm Beach, and Broward counties, Florida with that in Broward County persisting and becoming well established (Hammer, 2001). The recently discovered colony of S. paludicola in southwestern Sarasota County (Huegel & McConnell, 2008) is unquestionably indigenous and presents a significant range extension and a wealth of material for study. Based upon the several consistent differences between S. paludicola and S. lanceolata, the

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taxon is presented here at the species level and represents a near-endemic species for both the United States and Florida. Luer treated the taxon in the genus Spiranthes which was much the trend at that time. After Garay‘s (1982) revision of the Spiranthinae and reinstatement of the genus Sacoila, Sauleda, Hansen, and Wunderlin (1984) transferred the variety to the genus Sacoila. Luer noted most of the differences in his original publication (Luer, 1971) and reinforced them in Native Orchids of Florida (1972). Catling (1987) continued this work noting the breeding differences between S. paludicola and S. lanceolata. Although S. lanceolata is widespread and forms very distinctive clones throughout its range in the West Indies, Central and northern South America, plants have not been previously specifically noted that conform to any of the criteria for S. paludicola until specimens from western Cuba provided by Ackerman were examined. Luer‘s comment that S. paludicola would probably be found further south in the West Indies is supported by these collections from western Cuba. There are three additional orchid taxa that have a similar range limited to southern Florida and western Cuba are Galeandra bicarinata G. Romero & P.M. Brown, Dendrophylax lindenii (Lindley) Bentham, and Epidendrum floridense Hágsater. The critical differences between Sacoila lanceolata and S. paludicola are summarized below. Data for S. lanceolata is primarily from Florida material and for both is based upon work by Brown and Catling in FNA, (2002) with additional observations and recently collected data.
S. lanceolata absent at flowering time petiole indistinct to 65 cm tall glandular pubescent stout, to 8-11 mm clasping blending in with the stem 4-18 cm green to brown 9-50, brick red to terra cotta* tubular, apices distinctly spreading; apomictic 19–23 × 7–9 mm 23–31 × 3–8 mm 4–6 mm 21–26 × 6–8 mm lip pale red or pinkish white to white, 23–31 × 8–11 mm; margin slightly undulate 9–10 mm extending back 11-12 mm 8–9 mm 4–5 mm 10–20 mm 4–5 mm 14 × 8 mm polyembryonic dry roadsides, fields, open pine flatwoods, usually in sunny areas late April—mid-June 195 S. paludicola persistent (usually) at flowering time petiole distinct, tapering from blade to 40 cm tall pubescent slender, 4-6 mm foliaceous and slightly spreading at the tips; paler in color contrasting with the stem 3-15 cm red 5-29, deep red* tubular; apices only slightly spread; selfpollinating 9–11 × 3.5–4.5 mm 12–14 × 33–42 mm 0.8–1.2 mm 9–10.5 × 2.2–2.8 mm red to pale red, 11–13 × 3.8–4.5 mm; margin smooth 6–7.5 mm extending back 3-4 mm 3–3.5 mm 2.9–3.1 mm 12–15 mm 2.3–2.8 mm 12 × 6 mm monoembryonic hydric hammocks - trailsides, old logs, and stumps; in shaded areas late January—early April

leaves plant scape stem bracts inflorescences floral bracts flowers dorsal sepal lateral sepals mentum petals lip column pollinaria viscidium ovary rostellum capsules seeds habitat flowering time *excluding color forms

Pemberton et al: ALIEN TERRESTRIALIN RCHID, EULOPHIA GRAMINEA,TATES Brown: THE GENUS SACOILA O THE CONTINENTAL UNITED S INVADES MIAMI

Fig. 7 Sacoila paludicola (Luer) P.M. Brown 196

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Fig. 9 Sacoila squamulosa (Kunth) Garay

Fig.8 Sacoila lanceolata (Aublet) Garay

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Plants of Sacoila lanceolata in the Caribbean, Central and South America may contain localized populations/clones with monoembryonic seeds and occasionally have plants with persistent leaves at flowering time With the exception of the size of the pedunculate ovary, which falls within that of S. lanceolata, all measurements of S. paludicola are noticably lesser than those of S. lanceolata. When observed in the field the obvious differences, apart from the persistent leaves, are those of size, floral and bract color, habitat, flowering time, and the stem bracts. The less obvious, but critical, trait is the seeds. Specimens for the Florida records are located at SEL, FLAS, AMES, BUS (FTG), NY and the Cuban records at UPRRP. A golden yellow-flowered form was found in Corkscrew (Audubon) Sanctuary, Collier Co. Florida (Brown, 2001) and requires a transfer. Sacoila paludicola (Luer) P.M. Brown forma aurea (P.M. Brown) P.M. Brown (Fig. 5) Basionym: Sacoila lanceolata (Aublet) Garay var. paludicola (Luer) Sauleda, Wunderlin & B. F. Hansen forma aurea P.M. Brown North American Native Orchid Journal 7(1): 95-96. 2001.

Sacoila squamulosa (Kunth) Garay, Botanical Museum Leaflet 28: 352. 1982. (Fig. 6, 9)

hoary beaked orchid Basionym: Neottia squamulosa Kunth in A. von Humboldt et al., Nov. Gen. Sp. 1: 266, plate 71. 1815; Synonyms: Sacoila lanceolata var. squamulosa (Kunth) Szlachetko; Spiranthes squamulosa (Kunth) León; Stenorrhynchos squamulosum (Kunth) Sprengel Herbs, terrestrial or lithophytic, 30–60(–75) cm. Leaves absent at flowering time, 2–8, shortpetiolate; blade elliptic to broadly lanceolate, 30 × 5 cm, apex acute to obtuse. Inflorescences 8– 18 cm; rachis scurfy with white dots; floral bracts red, lanceolate, 15–25 mm, exceeding ovary, scurfy. Flowers brick red, 15–50; dorsal sepal 15–18 × 5 mm; lateral sepals 24 × 3.5 mm, connate for 5–7 mm over basal portion of mentum, distinct portion of mentum 15–18 mm, about 2/3 length of ovary; petals 18–24 × 3–6 mm; lip brick red to pale red, 20–24 × 6 mm; column 11 mm from attachment of dorsal sepal to tip, foot extending back 11–12 mm; pollinaria 4–5 mm, viscidia 3–3.5 mm; pedicellate ovary 10–13 mm; rostellum 4–5 mm. Capsules 10–13 × 7 cm, scurfy. Seeds largely monoembryonic. Flowering April--May. Open or wooded, dry--moist including poorly drained pinelands on calcareous substrates; 0--30 m; Fla.; Mexico; West Indies; Central America; n South America. Sacoila squamulosa is widespread in tropical and subtropical America. Although often overlooked in Florida, it was included in the range of Stenorrhynchos squamulosum (Nir, 2000), and details were given for its inclusion in the Florida flora (Brown, 2000). That resulted in determining that a large stand of nonconforming Sacoila in Marion County was indeed this species. It is a tall and handsome plant that, upon close examination, has the unmistakable scurfy or hoary dots instead of the short pubescence of Sacoila lanceolata. This is one of the most recent additions to the orchid flora of Florida and the United States. Plants have been either overlooked or not well distinguished from the more common S. lanceolata. REFERENCES:
Brown, P. M. 1999. A striking new color form of Sacoila lanceolata. North American Native Orchid Journal 5: 169–73. _____. 2000. Recent taxonomic and distributional notes from Florida 8: Sacoila squamulosa verified for the United States. North American Native Orchid Journal 6(4): 333–34. _____. 2001. Recent taxonomic and distributional notes from Florida 9: Sacoila lanceolata var. paludicola forma aurea. North American Native Orchid Journal 7(1): 95-96. Brown, P.M. & P.M. Catling. 2002. Sacoila in Flora of North America volume 26 Orchidaceae. Catling, P.M. 1987. Notes on the breeding systems of Sacoila lanceolata (Aublet) Garay (Orchidaceae). Annals of the Missouri Botanical Garden. 74: 58–68. Folsom, S.N. 2000. The Saga Continues. North American Native Orchid Journal 6(4): 330-32. 198

Brown: HE GENUS SACOILA O THE C EULOPHIA GRAMINEA, INVADES Pemberton et al:TALIEN TERRESTRIAL INRCHID,ONTINENTAL UNITED STATES MIAMI Garay, L.A. 1980 (1982). A generic revision of the Spiranthinae. Botanical Museum Leaflets. [Harvard University] 28(4): 277-426. Hammer, R.L. 2001. A status report on the native and naturalized Orchidaceae of Collier, Miami-Dade, and Monroe Counties, Florida. North American Native Orchid Journal. 7(1): 3-84. _____. 2002. Everglades Wildflowers. Guilford, Conn : The Globe Pequot Press. Huegel, C.N. and K.K. McConnell. 2008. General Notes on the Ecology of Sacoila paludicola in Sarasota County, Florida. North American Native Orchid Journal 14(3): 193-204. Luer, C.A. 1971. A variety of Spiranthes lanceolata in Florida. Florida Orchidist 14: 19. Nir, M. 2000. Orchidaceae Antillanae. New York: DAG Publishing. Sauleda, R., R.P. Wunderlin, and B.F. Hansen. 1984. Transfers in Florida Orchids. Phytologia. 56: 308. Jim Ackerman, University of Puerto Rico, provided scans of a number of Sacoila specimens including those now identified as S. paludicola from western Cuba and Bruce Hansen (USF) provided literature reprints.

Fig. 10 Sacoila paludicola 30 March 2008 Sarasota Co., FL

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GENERAL NOTES ON THE ECOLOGY OF SACOILA PALUDICOLA, SARASOTA COUNTY, FLORIDA
Craig N. Huegel, Ph.D. & Kathleen K. McConnell
INTRODUCTION

A previously unknown population of approximately 200 plants of the leafy beaked ladies‘-tresses, Sacoila paludicola (Luer) P.M. Brown, was recently discovered and confirmed by the authors in Sarasota County, Florida. This discovery represents a range extension in Florida of approximately 90 linear miles from the previous known locations and a new county record for this species. It also represents a significant increase in the known population of this state-listed threatened, terrestrial orchid. A description of the site and some preliminary observations on the distribution and ecology of this population is provided below.
SITE CONDITIONS

The exact location of this population is not being provided at this time to protect it from the potential impact of collectors, although the site is well protected, routinely managed, and unlikely to be developed. The population occurs in an area less than two acres in size. Although extensive searches were conducted elsewhere in the region, its restricted occurrence to this area is well defined by ecological conditions not present elsewhere on the property. Based on published written descriptions (e.g. Hammer, 2001; Hammer, 2002) and personal conversations (Mike Owen, pers. comm.), these conditions are very similar to those found for previously known populations from south Florida. Sacoila paludicola is restricted on this site to a region of hydric hammock that rarely floods (Figure 1). The forest community is characterized by a nearly closed canopy (>60 %) dominated by laurel oak (Quercus laurifolia) and cabbage palm (Sabal palmetto), although strangler fig (Ficus aurea), live oak (Quercus virginiana), sweet bay (Magnolia virginiana), swamp bay (Persea palustris), dahoon holly (Ilex cassine), and red mulberry (Morus rubra) are scattered throughout. Various woody shrubs comprise the mid-canopy, especially saw palmetto (Serenoa repens), wild coffee (Psychotria nervosa and P. sulzneri), and American beautyberry (Callicarpa americana). The understory is dominated by ferns, especially swamp fern (Blechnum serrulatum), marsh fern (Thelypteris palustris), and whisk fern (Psilotum nudum). In addition to S. paludicola, two other terrestrial orchids are common; the tooth-petal and long-horn false rein orchids (Habenaria odontopetala and H. quinqueseta, respectively). The soils are characterized as Floridana and Gator Soils; Depressional. These hydric soil types contain high organic concentrations, often consisting of peaty muck or mucky mineral components from the surface to a depth of around 14 inches. They are either saturated near the surface, or inundated for six to nine months each year. Soil permeability is rapid in the surface layer.
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Fig. 1 Hydric Hammock habitat where S. paludicola was found within the study site.

Unlike many floristically similar sites in south Florida, no evidence of feral hogs (Sus scrofa) was detected and they have not been present for at least the past decade according to the current land manager at the site. Some evidence of groundcover disturbance, caused by the ninebanded armadillo (Dasypus novemcinctus), was noticeable, but the characteristic large-scale rooting and vegetative destruction caused by hogs has not altered this forest understory.
RESULTS

The authors mapped and monitored individual Sacoila paludicola specimens February-May 2008 to gather preliminary information in an effort to better understand the distribution, ecological requirements, and general ecology of this species. During this effort, each plant was mapped and given a unique number for data collection purposes. One of these plants was collected as an herbarium specimen after flowering was initiated as was the above-ground portions of a second individual. It is the intent of the authors to monitor the individually marked plants in future years as part of a long-term ecological study. The distribution of Sacoila paludicola was not uniform throughout the hydric hammock forest or within the region of the forest where it occurred. Although detailed community structure measurements have not been collected, its distribution was noticeably restricted to the
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more open patches where saw palmetto, in particular, was less dominant. Figure 2 shows the non-random distributional pattern of this orchid within a portion of the study site and provides a visual reference to the type of habitat within this forest where it was most commonly found. For the most part, S. paludicola was more abundant along the edges of unimproved walking trails and within patches where the understory was less dominated by woody species. In these areas, the control of Brazilian pepper (Schinus terebinthifolius) has also served to maintain a greater degree of openness within the mid canopy than in other regions of the property. The land manager has made a concerted effort to remove the debris by hand from this nuisance plant control program rather than piling and leaving the material on-site. Although the site burned extensively during the summer of 1989, the effects of this fire seem to have been rather uniform throughout the hammock and do not seem to explain the distributional pattern of S. paludicola. Any subtle changes that might have been present immediately after the fire would likely have been lost during the intervening 19 years.

Fig. 2 Distribution pattern of S. paludicola within a portion of the hydric hammock forest. Orange flags mark individual plants.

Soil conditions also seem to be a major influence on the distribution of Sacoila paludicola within the project site. Although the high organic soils are uniformly present throughout the hydric hammock where this species occurs, the hydrology is more variable. The region occupied by this species does not seem to be inundated for extended periods, but seems to remain nearly saturated during most months. This relatively consistent hydrology seems significant to the distribution of this plant within the project site. Areas of extended or reduced hydrology from those occupied by S. paludicola (as evidenced by changes in the understory flora), do not seem capable of supporting this species.

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The number of leaves produced per plant was variable (range 1-5, mean 1.87 leaves, median 2 leaves) based on data collected from 172 individual plants with leaves remaining at the time of sampling (21 March – 3 April 2008). A total of 15 plants did not have leaves present during the sampling period. Data specific to leaves was not recorded for the remaining 13 individuals in our overall sample. The length of leaves was not accurately measured, but their general condition during flowering was assessed. Leaves that were brown or black were considered dead and were so noted. Leaves that were mostly green, but noticeably yellowing and showing signs of senescence were distinguished from leaves that were green and showing no signs of aging. Although these classifications are somewhat subjective, the characteristics used to differentiate them were obvious for most plants observed in the field. Figures 3-5 show leaves within the various classifications of leaf condition used in this analysis. Plants that had leaves that were not clearly differentiable between green and yellowing were not included in the analysis. Most of the 141 plants included in this analysis (n=96, 68.1%) had leaves that exhibited obvious signs of senescence. Three additional plants had leaves that were present, but obviously dead. The remaining 42 plants (29.8%) had leaves that were green and did not exhibit signs that they were beginning to senesce by 3 April.

Fig. 3 Flowering S. paludicola with ―dead‖ leaves.

Fig. 4 Flowering S. paludicola with ―yellowing‖ leaves, showing clear evidence of senescence.

Fig. 5. Non-flowering S. paludicola with ―green‖ leaves, showing no obvious sign of senescence.

Leaves were generally absent or dead by 8 May when flowering was completed and seed capsules were dehiscing. Of the 173 individual plants for which data was collected, 93 were completely devoid of leaves and 35 plants had at least one dead leaf still visible. A total of two plants still retained leaves that were green and showed no outward sign of senescence. The
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remaining 43 plants had at least one leaf that was noticeably yellowing. Plants did not lose their leaves at the same time, however. Nearly half (53%) of the plants that retained yellowing or green leaves on 8 May had lost at least one of their leaves that had been present on 3 April. In fact, both plants that retained a green leaf on 8 May had lost their other leaf between 3 April and this date. Plants do not remain leafless for long once their leaves are lost. Of the 173 individual plants reexamined on 8 May 2008, 31 had begun to leaf out once more. Figures 6 and 7 show two typical plants producing new leaves at this time; approximately one month after the completion of flowering and when most plants still retained at least one leaf seemingly capable of photosynthesis. A Chi-square analysis was performed on this data in an effort to determine if there was a relationship between early leaf production and the flowering status of individual plants. Plants that produced a flowering stalk in 2008 were significantly less likely to be among the 32 earliest plants to produce new leaves than plants that did not flower (X2 = 14.16, n = 4, p< 0.01). Only two plants that flowered in 2008 were found to be sending up new leaves on 8 May. In addition, two new plants not found during our initial surveys on 21 March 2008, seemingly because they had lost their leaves by this date, were discovered on 8 May and were marked for future study. This result suggests that other non-flowering plants were leafless by 21 March and were undetectable during the 2008 study period.

Fig. 6 Early leaf emergence of leafless S. paludicola, 8 May 2008. attachment.

Fig. 7 Early leaf emergence of S. paludicola with one ―yellowing‖ leaf. New leaf is barely evident at point of leaf

Most individuals did not flower during 2008. Of the 198 plants for which flowering data was collected, a total of 73 (36.9%) produced a flowering stalk. This result is biased, however, as it includes individuals that were either seedlings or likely seedlings based on their small leaf size. When individuals with noticeably small leaves were removed from the analysis (33 total), the percentage of individual plants that flowered increased to approximately 44.2%. This number is likely a more accurate estimate of the percentage of mature individuals that bloomed. Flowering in 2008 was quite synchronous and occurred during a four-week period between mid-March and mid-April. All flowers were a uniform scarlet red in color; figures 8 and 9 show typical flowering. Flower stalks were well-developed during a 24 February site visit, but the flower buds were immature and no open flowers were evident. Flowering was evident on a return field visit 15 March. Although detailed field data was not collected on this date, very few plants were in bloom on 15 March and those with open flowers generally had only one or two blooms at the lowest end of the flowering stalk. Flowering was approximately at its peak on 30
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March. All, but one, of the 73 plants that produced a flowering stalk were in bloom and all flowering stalks had at least one bud that had not yet opened. By 3 April, flowering was nearly completed. Of 57 individual plants for which data was collected on this date, 18 (31.6%) had completely finished blooming and another six individuals (10.5%) had open flowers, but no unopened buds. Only eight individuals (14.0%) had a total of more open flowers plus unopened flower buds than flowers that were past blooming. No flowering was evident on a 20 April site visit.

Fig. 8 Flower stalk of S. paludicola, 21 March 2008. Lower buds nearly ready to open.

Fig. 9 Flower stalk of S. paludicola, 30 March 2008. Lower flowers have finished, middle buds are open, and upper buds have not yet opened.

The number of flowers produced per individual plant was variable (range 5-25). Based on the sample of paludicola, 30 March 2008. Lower flowers have finished, flowers per open, and stalk was Fig. 9Flower stalk of S. 57 blooming individuals, the mean number ofmiddle buds arebloomingupper buds 12.6. yet opened. have not This figure, however, was not representative of the overall population as 35 individuals (61.4%) produced fewer flowers than the population mean. Because of this, the median number of flowers per individual plant (11) may provide a better estimate of the average number of flowers per individual. Nearly every flower/flower bud produced a seed capsule. Figures 10 and 11 show the near 100% development of seed capsules from flowers that opened. Of the 57 individual plants for which data was collected, only one plant that produced a mature flower stalk failed to flower. The cause for the loss of this flower stalk could not be determined, but was likely due to herbivory. Of the remaining 56 flowering plants, 53 (94.6%) produced a ripened seed capsule for

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every flower bud. Of the remaining three plants, two plants produced ten ripened seed capsules out of 11 flowers and the remaining plant produced two capsules from a total of five flowers.

Fig. 10 Flower stalk of S. paludicola, 3 April 2008. Fig. 11 Flower stalk of S. paludicola, 8 May 2008. Lower flowers have developed into ripening seed capsules. Seed capsules have dehisced and some seed is Only the uppermost flower buds remain to still open. evident on flower stalk, capsules, and on spider web.

The relationship between leaves and flowering is not clear and warrants additional data collection. As stated above, 15 plants flowered after their leaves had died and were no longer visible. For the remaining 51 flowering plants for which data was collected in 2008, there was a positive relationship between the number of leaves produced and flowering (X2 =11.85, n=5, p<0.05). Plants having a larger number of leaves than average produced a flowering stalk in greater-than-expected ratios than plants that had fewer leaves. Every plant that had four (n=4) and five (n=1) leaves produced a flower stalk as did 13 of 26 mature plants that produced three leaves. Plants having one or two leaves produced fewer flower stalks (12 of 51 and 21 of 62 plants, respectively). Plants with seemingly healthy leaves at the peak time of blooming tended to flower less frequently than those with leaves that were obviously yellowing. Although the majority of plants for which data was recorded were losing their leaves or had already lost them at the time of flowering, a slightly greater-than-expected (X2=7.04, n=3, p< 0.07) number of plants having leaves that were obviously dying had flowering stalks. Plants that lost their leaves, but did not flower, could not be detected by the authors and were not included in this analysis.

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DISCUSSION

Until recently (Brown 2008), the leafy beaked ladies-tresses has been regarded as a variety of the more common and widely distributed leafless beaked ladies‘-tresses, Sacoila lanceolata. Though S. lanceolata has been described from nearly every county in peninsular Florida, from Walton County within the Florida panhandle, and from the Caribbean, Central and South America as far south as Uruguay, S. paludicola has been known to occur in only a few counties in extreme south Florida. A recent review of herbarium specimens collected by J.D. Ackerman, M.A. Ruiz, and J.L. Bocourt, and reported by Brown (2008), from broadleaf forested areas within the Pinar del Rio Provence in western Cuba confirm its presence outside of Florida. Other Caribbean records of Sacoila spp., however, have not been verified as S. paludicola. Populations found in the general vicinity of the type location described by Luer (1971) from observations first made January 1965 within the Fakahatchee Strand State Preserve and later from nearby Corkscrew Swamp in Collier County are naturally occurring. It has not been described as occurring within the nearby Florida Panther National Wildlife Refuge, (Stewart and Richardson, 2008) and there is question that other populations (i.e. Miami-Dade, Broward and Palm Beach Counties) were purposely introduced (Hammer, 2001, P.M. Brown, personal communication). It is certain that Frank Craighead made several attempts to introduce it from the Fakahatchee Strand to areas within Everglades National Park during the mid-1900s, that one population continues to persist in Grossman Hammock inside the park boundary, and that several other populations persisted for years (and possibly disseminated seed) before finally disappearing (Hammer, 2001). Hammer (2001) also cites anecdotal information questioning the natural occurrence of the colony recently found in Broward County. As such, S. paludicola has one of the most-restricted natural ranges of any of Florida‘s orchid species. Sacoila paludicola has not been well studied and, for the most part, little more than anecdotal comments have been published in the literature. Differences between the two species, however, have been well noted and are easily recognizable. Although Luer (1971) considered the differences between these two species ―minor‖, he described numerous characteristics that differentiated them, including habitat preference; leaf shape, persistence and ―glossiness‖; flower color and size; and blooming season. He also noted that these differences were not the result of growing-condition influences; that plants of each variety maintained their unique differences even when grown in pots next to each other and given the same culture. Despite this, he attributed their differences to natural variation and conjectured that S. paludicola might be the more widespread form in ―tropical America‖, seemingly present in shady moist forested understories similar to the conditions found in Fakahatchee Strand. Specimens of S. paludicola collected in western Cuba were found growing in conditions seemingly similar to those found for this species in Florida. This has not been the habitat type, however, reported for specimens of the genus collected in Puerto Rico where it was found on roadsides and pastures (J.D. Ackerman, pers. comm., reported in Brown 2008) or in Brazil where it is reported to occur along roadsides from sea level to elevations of approximately 4,000 feet and not within tropical portions of the country (Francisco Miranda, pers. comm.). The discovery of a second significant natural Florida population of S. paludicola in Sarasota County in 2008 and the verification of a population in western Cuba, seem to demonstrate that the population centered in and around the Fakahatchee Strand is not an aberrant form of S. lanceolata adapted to localized environmental conditions. The habitat requirements and morphological differences noted by Luer (1971) are identical for both the Sarasota County and extreme south Florida populations and seem to be shared by the herbarium specimens collected in western Cuba.
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Sacoila paludicola seems to be narrowly restricted by habitat type. Unlike the morecommon S. lanceolata, it is not found in open sunny habitats with varying soils and soil-moisture conditions. In Sarasota County, all individuals of S. paludicola were found in hydric hammock forest characterized by organic soils and relatively stable hydrology. Within these areas, plants were largely confined to the edges of unimproved trails and other patches where the understory was not dominated by saw palmetto and other woody species. Such conditions are very similar to areas within the Fakahatchee Strand State Preserve, where most individual plants are confined to historic tram trails inside the swamp forest that flood very infrequently (Mike Owen, pers. comm.). Other characteristics reported by Luer (1971) and by others to distinguish Sacoila paludicola from S. lanceolata are also evident in the Sarasota County population. Figures 12 and 13 show the obvious differences between the flowers of the two related species. Flower coloration in the Sarasota population of S. paludicola was the same scarlet color as has been previously documented and not the pink-salmon color typical of most S. lanceolata. Additionally, the variation in color, sometimes evident in S. lanceolata was not evident in the Sarasota population. P.M. Brown (2001) documented one individual S. paludicola in Corkscrew Swamp Sanctuary with a yellow-colored flower, forma aurea, but this plant has since disappeared (P.M. Brown, pers. comm.) and no others have been reported. In addition, no scarlet-colored forms of S. lanceolata

Fig. 12 Flower stalk of S. lanceolata, 10 May 2008. Specimen from southern Hardee Co., FL. Flower color and structure differ markedly from that of S. paludicola. . 208

Fig. 13 Close-up of S. paludicola flowers in contrast to S. lanceolata, shown opposite.

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have been reported, similar to the color found almost universally in S. paludicola. Various morphological differences between the two species are also evident in the photographs. The flowering period of the Sarasota population of Sacoila paludicola is nearly identical to that found in the Fakahatchee Strand, based on nearly a decade of unpublished observations recorded by Preserve biologists and shared with the authors (Mike Owen, pers. comm.). It also seems to closely correspond to the blooming season of herbarium specimens of S. paludicola collected while in flower in western Cuba. S. paludicola in Florida seems to have a narrow flowering season that occurs between mid-March and mid-April. This differs markedly from accounts reported in the literature for S. lanceolata in Sarasota County and further south, within the current range of S. paludicola. Literature reports (Hammer, 2002; Brown and Folsom, 2005; Stewart and Richardson, 2008) and observations of the authors suggest that S. lanceolata does not initiate blooming until late April and that the peak of blooming occurs sometime in May. Sacoila paludicola has adopted a different pollination strategy than that adopted by S. lanceolata. Our observations seem to confirm the observations of Catling (1987) that S. paludicola is self-pollinating. Despite numerous hours of field observation by the authors, no evidence of pollination by traditional pollinators of this species (i.e. bees and hummingbirds) was recorded. On several occasions, open flowers were visited by Gulf fritillary butterflies (Agraulis vanillae) (Figure 14), but these visits were infrequent and are not likely to account for the nearly 100 percent development of ripe seed capsules from open flowers within the study area. Similar to that reported by Catling (1987), individual flowers were short-lived, but no effort was made by the authors to observe whether pollinaria were removed or not. Although Catling (1987) demonstrated that pollination was necessary for seed development in Sacoila lanceolata in both Ontario, Canada and in central Guyana, south Florida populations (including the latitude of the Sarasota study area) of S. lanceolata have adopted a pollinator-independent strategy. This strategy, however, is different from that of S. paludicola; being apomictic instead of self-pollinating. Catling (1987) postulated that this trait developed uniquely in south Florida as a response to a lack of hummingbirds during the later May-June blooming season. Hummingbirds are common residents of both Ontario and Guyana during the blooming season of S. lanceolata, but are uncommon in south Florida at this time of year (Sarasota Audubon Society, 2008). Although the presence or absence of hummingbirds may adequately explain the development of a pollinator-independent strategy in south Florida populations of Sacoila lanceolata, it is not likely the sole selection pressure for S. paludicola. Though ruby-throated hummingbirds (Archilochus colubris) are generally absent from Collier and Miami-Dade Counties during the flowering period of S. lanceolata (Austin 1975), they are not uncommon migrants in extreme south Florida one month earlier at the time when S. paludicola is in flower. The universal development of a pollinator-independent reproductive strategy in S. paludicola is more likely a response to the selection pressure required by its different habitat requirements. In the open habitats along roadsides and pasturelands where S. lanceolata is most frequently observed throughout its geographical range, pollination by hummingbirds is easily facilitated. Rubythroated hummingbirds, however, are less likely to occur in the shady forested habitats where S. paludicola occurs, even though this species is present in the region during late March and April. The development of a universal strategy for self-pollination would be extremely advantageous in this setting.
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In addition to the differences between the two Sacoila species in blooming season and pollination strategy, S. paludicola exhibits a different strategy in maintaining its leaves for longer periods of time and closer to blooming. As has sometimes been mistakenly reported (Hammer 2002), S. paludicola does not always have leaves at the time of flowering, but typically maintains them to the time of flower stalk emergence. This characteristic is in sharp contrast to S. lanceolata which typically loses its leaves 30-90 days prior to the emergence of its flowering stalk. Although a majority of the plants had leaves that exhibited obvious signs of senescence at the time of blooming, only a very small minority (n=18) of the 187 plants for which this data was collected had already lost their leaves or had dead leaves that were still present. In addition, nearly onequarter of the plants (n= 42) maintained healthy-looking leaves past the peak of blooming on 3 April. The maintenance of leaves is likely another adaptation to life within shady forested habitats. While S. lanceolata is found most commonly growing in open, sunny habitats where abundant energy required for blooming can be acquired and stored in a shorter period of time, S. paludicola is resident to habitats where far less solar energy reaches its leaves. Under these conditions, maintaining leaves for a longer period may be necessary to store sufficient energy to allow for the development of a flowering stalk. Our data suggests that the total surface area of leaves for blooming plants was greater than that for non-blooming ones. Long-term study of individually marked plants and accurate measurements of leaf area may verify this result. The differences between Sacoila paludicola and S. lanceolata are great and seem to be the result of long-term adaptation to the vastly different habitat conditions exploited by each species. Our observations suggest that the population discovered in Sarasota County exhibits similar morphological characteristics and ecological requirements to previously known populations in extreme south Florida; characteristics and requirements that are quite dissimilar to populations of S. lanceolata within the same geographical area as S. paludicola and elsewhere. Although further study is warranted, this stability within disparate populations of each individual species throughout its known range and the stark differences between the two species irrespective of geographic range firmly suggests that these are not varieties of the same species, but separate ones.
ACKNOWLEDGEMENTS

The authors wish to thank the administration and staff of the study site for their support and assistance throughout all aspects of this project. We are particularly indebted to the site manager, S.H.K., for the enthusiasm and interest provided us. The fieldwork would not have been possible without the support of the authors‘ employer, Biological Research Associates, Inc. which provided time and materials throughout the study to complete the fieldwork needed for this paper and the philosophical support necessary for all other aspects. Our research was greatly assisted by the involvement of Paul Martin Brown, who provided significant insight and direction to our work and Wesley E. Higgins, Ph.D., Head of Systematics, Marie Selby Botanical Gardens. We also wish to thank the individuals that provided supporting information for this paper by personal communication; Mike Owen, Fakahatchee Strand State Preserve; James D. Ackerman, Ph.D., Florida State University; and Francisco Miranda, author and orchidologist, who has traveled extensively studying orchids in his native Brazil. All images used in the article were taken by the senior author, except Figure 1 which was taken by the junior author.
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LITERATURE CITED

Austin, D.F. 1975. Bird flowers in the eastern United States. Florida Scientist 38: 1-12. Brown, P.M. 2001. Recent taxonomic and distributional notes from Florida 9. North American Native Orchid Journal 7(1): 91-98. _____. 2008. The Genus Sacoila in the Continental United States. North American Native Orchid Journal 14(3): 192-199. Brown, P.M. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and expanded edition. Gainesville: University Press of Florida. Catling, P.M. 1987. Notes on the breeding systems of Sacoila lanceolata (Aublet) Garay (Orchidaceae). Annals of the Missouri Botanical Garden. 74(1): 58-68. Hammer, R.L. 2001. A status report on the native and naturalized Orchidaceae of Collier, MiamiDade, and Monroe Counties, Florida. North American Native Orchid Journal. 7(1): 3-84. _____. 2002. Everglades Wildflowers. Guilford, Conn : The Globe Pequot Press. Luer, C.A. 1971. A variety of Spiranthes lanceolata in Florida. Florida Orchidist. 14: 19. Sarasota Audubon Society. Home Page. 7 March 2008. Birding in Sarasota. <http://www.sarasotaaudubon.org/birding%20in%20sarasota.htm>. Stewart, S.L. and L.W. Richardson. 2008. Orchid flora of the Florida Panther National Wildlife Refuge. North American Native Orchid Journal. 14(2): 70-104. Craig N. Huegel, Ph.D. Biological Research Associates, Inc., a Division of ENTRIX, Inc. Sarasota, FL chuegel@biologicalresearch.com Kathleen K. McConnell Birkitt Environmental Services, Inc. Tampa, FL kmcconnell@birkitt.com

Fig. 14 Gulf fritillary butterfly (Agraulis vanillae) visiting the open flowers of S. paludicola. Butterfly visitation was rarely observed and does not seem to explain the near-100% pollination rates observed in 2008.

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with references to extant sites in New York, Pennsylvania and Nova Scotia
Paul Martin Brown This paper is an adaptation of a graduate thesis submitted to the University of MassachusettsDartmouth in June 1997. It is being presented here at the request of a number of Journal readers. NOTE: In the ten years since this paper was written succession and, in a few instances road work has taken its toll on many of the Spiranthes casei sites mentioned below. Natural succession has been the primary culprit and unless the roadsides were regularly mown, they have grown up and the Spiranthes are no longer present, at least as flowering plants. In almost all instances additional sites may be found within a short distance of the original sites documented herein. This is a classic case of habitathunting. For that reason I am reluctant to state that specific sites are no longer extant in 2008 until the general areas are well searched. Recently S. casei has also been seen in New Brunswick and near Scranton, Pennsylvania. The New Brunswick report is especially important as it may open the possibility of additional sites in northeastern Maine. In 1995 many areas along Route 1 in New Brunswick were examined but no S. casei was found and the same for numerous Aroostook Co., Maine areas with typical habitat. But that may all have changed by now. If you are so disposed to get out there and hunt for them late August into early September is the time. If you are successful please let me know so I can update my records. Good hunting! PMB ABSTRACT The genus Spiranthes L.C. Richard, the ladies‘-tresses orchids, is a small but complex genus of morphologically similar terrestrial orchids found primarily in North America. Much confusion has existed as to the delimitation of the species and their accurate distribution because of the similarity of species. Spiranthes casei Catling and Cruise var. casei was described in 1973 and corresponds to what was previously known as the ―northern (Spiranthes) vernalis.‖ A review of the taxonomic history of the species as well as a comparison of the characteristics of the similar species will also be discussed. Since its description the complete range of the species has not been well understood due to the lack of herbarium material and extensive field work. This present work will attempt to
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carefully and accurately define the range of the species in New England, based on field work from 1989-1996. Extensive references will also be made to additional sites and habitats in New York, Pennsylvania and Nova Scotia. The concept of genus communities and presence of nothospecies and hybrids will also be discussed in relation to several of the major sites for Spiranthes casei. INTRODUCTION The genus Spiranthes L. C. Richard, the ladies‘-tresses orchids, is represented in New England by the presence of eight species, one variety and three nothospecies as summarized below: Spiranthes casei Catling & Cruise var. casei Spiranthes cernua (L.) L.C. Richard Spiranthes lacera Rafinesque var. lacera Spiranthes lacera Rafinesque var. gracilis (Bigelow) Luer Spiranthes lucida (H.H. Eaton) Ames Spiranthes ochroleuca (Rydberg) Rydberg Spiranthes romanzoffiana Chamisso Spiranthes tuberosa Rafinesque Spiranthes vernalis Engelmann & Gray Spiranthes ×borealis P.M. Brown Spiranthes ×eamseii P.M. Brown Spiranthes ×intermedia Ames Spiranthes ×simpsonii Catling & Sheviak. The expedient identification of taxa within the genus is often complicated by both cryptic hybrids and, especially within the Spiranthes cernua complex, a number of geographic races or ecotypes that often represent extensive gene flow from other species (Homoya 1994; Sheviak 1982, 1991). Northern New England (defined as the counties of Chittenden, Washington, Caledonia, Essex, Grand Isle, Franklin, Lamoille, and Orleans in Vermont; Grafton, Carroll, and Coos in New Hampshire; and Oxford, Franklin, Somerset, Penobscot, Piscataquis, Aroostook, Washington, and Hancock in Maine) has six of these species and two nothospecies present, as well as two locally occurring hybrids. They are: Spiranthes casei Catling and Cruise var. casei Spiranthes cernua (L.) L.C. Richard Spiranthes lacera Rafinesque var. lacera Spiranthes lucida (H.H. Eaton) Ames Spiranthes ochroleuca (Rydberg) Rydberg Spiranthes romanzoffiana Chamisso Spiranthes ×borealis P. M. Brown Spiranthes ×simpsonii Catling & Sheviak Prior to 1974 Spiranthes vernalis Engelmann & Gray was also listed for northeastern Vermont and northern New Hampshire (Ames 1905, 1921, 1924; Pease 1964; Seymour, 1969). TAXONOMIC HISTORY OF SPIRANTHES CASEI Catling & Cruise The ladies‘-tresses orchid now known as Spiranthes casei Catling & Cruise was originally described as occurring from Ontario, Quebec, Nova Scotia, Michigan, Vermont, and New
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Hampshire (Catling & Cruise 1974). At the time of its original publication, specimens had been previously labeled as Spiranthes vernalis or, in a few instances, Spiranthes cernua var. ochroleuca. (Catling & Cruise 1974, p. 531; Mousley 1924, 1941, 1942). These plants were often referred to as the ―northern vernalis‖ (Case 1964, p. 86-87; Case 1987, p. 168-169; Correll 1950, p. 227; Voss 1972, p. 461-463). Until recently, field verification for this ―northern vernalis” had been minimal. New England specimens were cited from Sutton, Orleans County, Vermont Mrs. Bruche s.n. , 1899 (NEBC/AMES) and Colebrook, Coos County, New Hampshire A. S. Pease 29802, 1942 (NEBC/AMES). Further herbarium searches have since revealed specimens from additional towns near both of those collections. In 1975 Catling collected Spiranthes casei var. casei near Rangely, Maine. In 1980 Frankie Brackley (1985, pp. 74, 108) found a few plants of Spiranthes casei var. casei in an old field near New Hampshire Route 145 in Colebrook, New Hampshire. In 1989, while preparing for a major work on the native orchids of New England (Brown 1993, 1997) two populations of Spiranthes casei var. casei were discovered along New Hampshire Route 145. The first, in Colebrook, was on a west-facing slope with a variety of woody and herbaceous material, primarily Danthonia spicata, Solidago nemoralis, Thuja occidentalis, and Picea rubra. Spiranthes ochroleuca was also present on this slope. The slope is often mowed annually in late summer, decapitating many of the plants. In 1989 there were thirty-four flowering spikes at this station. Further north, on the same road, in Stewartstown Hollow, New Hampshire, a dry shaley slope with more than forty flowering plants was found. Except for the fact that the slope faced east, the companion plants were the same, although much sparser. Earlier in the season, Spiranthes romanzoffiana was flowering here. This site is particularly interesting because fourteen taxa of orchids can be found in less than thirty-five meters of roadside (Brown 1996)! In 1990 five additional sites were found in and near Colebrook, as well as one site on New Hampshire Route 16 in Dummer. Three of the additional sites were found on New Hampshire Route 145 and were located on very predictable slopes. The number of flowering plants varied from five to thirty, plus many more that had been mowed down. Two small sites were found adjacent to a gravel pit on New Hampshire Route 26 east of Colebrook. Careful examination of the soil and vegetation at all of these sites allowed a group of students led by the author to then go to Vermont and search for appropriate habitat in the vicinity of the historical locations in that state. We centered our search around the Lake Willoughby region in Orleans County. Most of the open habitat there proved too wet, although it supported large populations of Spiranthes cernua, Spiranthes romanzoffiana and, occasionally in the drier areas, Spiranthes ochroleuca. The first suitable slope that was found had Spiranthes casei var. casei on it. Only five plants in a very precarious cattle-traveled area were flowering. But, as in New Hampshire, it was shaley, with the Danthonia and Solidago present. As soon as other grasses and herbs predominated, Spiranthes ochroleuca was present. Further searching led to three other roadside slopes with Spiranthes casei var. casei. An additional population was seen in the center strip of US Interstate 91 just south of the Lyndonville, Vermont exit. Traffic and time did not permit exploration of this or additional possible sites on the Interstate. Suitable habitat extended for about one mile south of the exit. On September 7, 1990 Catling‘s site northeast of Rangeley, Franklin County, Maine was relocated. Extensive, suitable habitat exists in this locality. Only a few plants were found due to excessively heavy rains that autumn. Careful searching of the immediate area in subsequent years has revealed only a few more plants.
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During intensive field work in 1995 and 1996 ten additional sites were located and have been carefully documented. All observations of flowering plants were made between 25 August and 10 September each year. The following is based upon the original description of Catling and Cruise (1974). Additional notes are intended to supplement the description. Spiranthes casei Catling and Cruise var. casei Rhodora 76(808): 526-536. 1974. Figure 1. Plants (13-)27-33(-43) cm tall at anthesis. Leaves glabrous, often glaucous, ascending up to 1/3 the length of the stem; lowest leaves, not persisting until anthesis, ovate-lanceolate, 1-2 cm wide, 7 cm long; upper leaves, persisting until and sometimes throughout flowering, oblanceolate or linear-lanceolate, 15 to 20 cm long, and usually less than 1 cm wide, reduced to bladeless sheaths below the inflorescence. Stem glabrous basally, becoming pubescent in the rachis with septate, reddish-glandular hairs 0.10-0.30 mm long. Spike usually 6-10 cm long, occasionally to 15 cm long, with the cream-colored flowers arranged loosely and often in a single spiral. Floral bracts ovate or ovate-lanceolate, long-acuminate, 7-12 mm long, more or less pubescent basally. Ovary 4-6 mm long at anthesis, obliquely swollen on the upper side. Pubescence on the surface of the ovary, dorsal sepal, and lateral sepals, similar to that of the rachis. Lateral sepals 5-7 mm long. Dorsal sepal and petals barely upcurved, 5-7 mm long, the dorsal petals papillose on the outer surface. The lip 6.0-6.5 mm long in the lower flowers, 5-6 mm long in the upper flowers. Basal calli of the lip stout, 0.8-1.0 mm long, incurved. The lower surface and central upper surface of the lip, and region of the calli short pubescent or papillose. The early senescence of the leaves which is noted here, and also by Case (1984), does not appear to apply throughout the range of the species. In northern New England, New York, and Pennsylvania, basal leaves are usually in good condition at the onset of anthesis. TYPE: CANADA: Ontario: MUSKOKA DISTRICT: ca. 9 km west of Bracebridge along highway 118 in Monck township, on dry hillside in full sun, Catling & Whiting (TRT 169205). REPRESENTATIVE SPECIMENS CANADA: NOVA SCOTIA: Shelburne Co.: Hope's Lot Barrens, Clyde River, 7 Sept. 1921, M. L. Fernald & B. Long (AMES 85418). ONTARIO: Frontenac Co: Salmon Lake, Frontenac Park, Bedford Twp., 23 Aug. 1968, R. Hainault & I. MacDonald (QK 97607). Simcoe Co.: lot 4, conc. 10, Innisfill Twp., 30 Aug. 1970, T.A. Reznicek (Simcoe Co. Museum 470) QUEBEC: Stanstead Co.: Hatley, 11 Sept. 1924, Mousley (QFA 42012, AMES 63699). UNITED STATES: MICHIGAN: Marquette Co.: summit of Huron Mountain, 1-9 Sept. 1916, C.K. Dodge (MICH). NEW HAMPSHIRE: Coos Co.: Colebrook, 21 Aug. 1942, A.S. Pease 29802 (NEBC). VERMONT: Orleans Co.: Sutton Rd., Willoughby, 13 Sept. 1899 Mrs. Bruche (NEBC). Current range extended to Wisconsin, Maine, New York, and Pennsylvania (Brown 1993, 1997; Chapman 1997; Hapeman 1996; Mitchell 1986). The specimen cited from Nova Scotia became the type for Spiranthes casei var. novaescotiae.

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Fig. 1 Spiranthes casei var. casei, Coos Co., NH
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SPIRANTHES CASEI VAR. CASEI SITES IN MAINE, NEW HAMPSHIRE, AND VERMONT

Map 1. General distribution of populations of Spiranthes casei var. casei in northern New England MAINE. Franklin County Site 1. Dallas Plantation; 6.75 km northeast of Rangeley, shaley roadside bank, P.M. Catling, 31 August 1975 (TRT) This same locality and adjacent shaley barrens were examined in 1990 and a few flowering plants, as well as several non-flowering plants, were found. Extensive suitable habitat exists in this area, although few plants have ever been located. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1991 - 1; 1992 - 3; 1993 - not field checked; 1994 - 1 - different area; 1995 - not field checked; 1996 - 0. Oxford County Site 2. Magalloway River and Maine Route 16; Located on a large, thin-soil, banking on the north side of Magalloway River Road. Other Spiranthes species typical of xeric habitats can be found here as well as additional species of orchids. Additional taxa of orchids noted: Spiranthes cernua, Spiranthes lacera var. lacera, Spiranthes romanzoffiana; Malaxis unifolia. Flowering plants of Spiranthes casei var. casei observed: 1996 - 3 plants in fruit.
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NEW HAMPSHIRE. Coos County Site 3. West Milan i This very small, but productive and reliable site, in West Milan is on the east side of New Hampshire Route 110, 0.75 km south of New Hampshire Route 110A. It was first found in 1993 and presented problems for the identification of several of the plants. Eventually these problematic individuals were described from this site as Spiranthes ×borealis P.M. Brown (Brown 1995). It has proven to be one of the most reliable sites for Spiranthes casei var. casei in the Northeast, producing several flowering plants each year. Additional taxa of orchids noted: Spiranthes lacera var. lacera, Spiranthes ochroleuca, Spiranthes ×borealis. Flowering plants of Spiranthes casei var. casei observed: 1993 - 13; 1994 - 12; 1995 - 14; 1996 - 31. Site 4. West Milan ii A single flowering plant of Spiranthes casei var. casei was found in 1996 on the west side of New Hampshire Route 110, 0.5 km south of New Hampshire Route 110A, on a raised bluff. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1996 - 1. Site 5. Dummer i This is an extensive Spiranthes site that stretches along the highway for more than 0.75 km on west side of New Hampshire Route 110, 0.5 km north of New Hampshire Route 110A. The plants are scattered along a rocky outcrop. Spiranthes casei var. casei is the least abundant Spiranthes at this site; four other species are found in larger numbers. Spiranthes cernua, Spiranthes ochroleuca and especially Spiranthes lacera var. lacera, which occurs in great numbers — exceeding one thousand flowering plants in 1996. One of the curious observations in 1996 of Spiranthes lacera var. lacera was the presence of four cleistogamous flowering plants. Additional orchid taxa noted: Spiranthes cernua, Spiranthes lacera var. lacera, Spiranthes ochroleuca. Flowering plants of Spiranthes casei var. casei observed: 1993 - 6; 1994 - 3; 1995 - 0; 1996 - 1. Site 6. Dummer ii A small sand bank habitat on the west side of New Hampshire Route 16 just south of the Cambridge/Dummer town line. Numerous plants of Spiranthes lacera var. lacera, Spiranthes cernua, and Spiranthes ochroleuca occur at this site. There have never been many plants of Spiranthes casei var. casei found here. What makes this site unusual is the sand substrate as opposed to the shaley substrate elsewhere in New England. In this respect it is similar to sites seen in northern New York State. On the east side of the road Spiranthes lacera var. lacera, Spiranthes romanzoffiana and the rare nothospecies Spiranthes ×simpsonii can be found as well as several other species of orchids. Additional orchid taxa noted: Spiranthes cernua, Spiranthes lacera var. lacera, Spiranthes ochroleuca, Spiranthes romanzoffiana, Spiranthes ×simpsonii; Gymnadeniopsis clavellata var. ophioglossoides, Platanthera huronensis. Flowering plants of Spiranthes casei var. casei observed: 1991- 1; 1992 - 0; 1993- 0; 1994 - 0; 1995 - 2; 1996 - 0. Site 7. Columbia Bridge First found in 1995, this excellent site is on the east side of US Route 3, 8.25 km south of Colebrook and 0.75 km north of Columbia Bridge Road, along the Connecticut River. The habitat is a very steep, rocky slope that is overgrown with small to medium clumps of deciduous and evergreen trees. The rocks are primarily schist and shale. The plants of Spiranthes casei var.
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casei grow in small pockets throughout the rocks. The roadside mowing is restricted to the lower eight feet of the slope so the plants that occur above that line are not damaged by the mowing blade. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1995 - 14; 1996 - 19. Sites 8/9. Kidderville a/b. This and the following site (site 9) are located adjacent to each other along New Hampshire Route 26 on the south side of the road just east of the Redwood Motel. Plants were observed only in 1990 at both sites, although the area has been searched each of the other years. Additional orchid taxa noted: Spiranthes cernua, Spiranthes ochroleuca; Epipactis helleborine. Site 8. Flowering plants of Spiranthes casei var. casei observed: 1990 - 3; 1991-1996 - 0. Site 9. Flowering plants of Spiranthes casei var. casei observed: 1990 - 1; 1991-1996 - 0. Site 10. Colebrook i This is the first of a series of sites along New Hampshire Route 145 north of Colebrook (Map 13). This area from Colebrook through Stewartstown appears to be the stronghold for Spiranthes casei var. casei in New England. Although several populations are small in size, it is a reliable area to see plants each year. Populations are subject to mowing and roadside work. 1996 was one of the very best years to observe Spiranthes species along this roadway as no mowing took place after August 1. Unfortunately those sites which were repeatedly mown for several consecutive years did not yield plants in 1996. Additional orchid taxa noted: Spiranthes cernua, Spiranthes ochroleuca. Flowering plants of Spiranthes casei var. casei observed: 1989 - 34; 1990 - 31; 1991 - 12; 1992 - 6; 1993 - 14; 1994 - 0; 1995 - 0; 1996 - 0. Sites 11/12. Colebrook: Caron i (upper pit); Caron ii (lower pit and seep) First discovered in 1996, these contiguous sites off of New Hampshire Route 145 occur in a large pit area with an excellent Spiranthes genus community. The habitat is varied from wet seeps to xeric shales, which, in turn, support three species of Spiranthes and one nothospecies. All species were in good condition and offered excellent opportunities for comparison. Additional orchid taxa noted: Spiranthes cernua, Spiranthes ochroleuca, Spiranthes ×borealis. Flowering plants of Spiranthes casei var. casei observed: 1996 - 13 at site 11, Caron i, Map 15; 41 at site 12, Caron ii. Site 13. Colebrook ii This site on New Hampshire Route 145, at pole 67, is along a small roadside edge and was first noted in 1996 with nine flowering plants of Spiranthes casei var. casei. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1996 - 9. Site 14. Stewartstown: Bishop Brook Road Located on New Hampshire Route 145 just north of Bishop Brook Road at pole number 97/67, this population produced many plants in the early 1990‘s but continued mowing appears to have eliminated the population. Additional orchid taxa noted: Spiranthes cernua; Epipactis helleborine, Platanthera lacera. Flowering plants of Spiranthes casei var. casei observed: 1990 - 21; 1991 - 14; 1992 - 0; 1993 - 12; 1994-1996 - 0 (heavily mown); 1996 - 0.
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Site 15. Stewartstown This site, along New Hampshire Route 145 at pole 141, is most unusual because it does not contain Spiranthes casei var. casei, but does have a large population of Spiranthes ×borealis as well as many Spiranthes ochroleuca. It is noted because it is the only site found with the nothospecies and lacking Spiranthes casei var. casei. This site is situated only 100 m north of one of the best sites for Spiranthes cernua and Spiranthes ochroleuca and numerous plants of the distinctive, single-ranked S. cernua. Additional orchid taxa noted: Spiranthes ochroleuca. Flowering plants observed of Spiranthes ×borealis: 1996 - 34. Site 16. Stewartstown Hollow This site on New Hampshire Route 145 at pole number 153 - 6/1 1/2, is similar to site 14 at Bishop Brook Road in that it has been mown repeatedly until few, if any, flowering plants have been observed for several years. The reappearance in 1995 of four plants is hopeful. Additional orchid taxa noted: Spiranthes cernua; Epipactis helleborine, Platanthera lacera. Flowering plants of Spiranthes casei var. casei observed: 1990 - 6; 1991 - 2; 1992 - 8; 1993-1994 - 0 (closely mown); 1995 - 4; 1996 - 0. Site 17. Creampoke i This site, and the following (site 18, Creampoke ii), were the two most significant sites to be studied. It was here that the best opportunity for making long-term observations was found in sites that are less disturbed by mowing. Creampoke i is located on New Hampshire Route 145 on an east-facing slope just south of Creampoke Road in Stewartstown Hollow. The habitat is a combination of mown, scrubby roadside and an open, xeric shale mound that is mown (or more appropriately brushed out) every ten years or so. It has only been heavily cut once since 1990. The plants of Spiranthes casei var. casei are well distributed throughout the site and reach their best development in the undisturbed, hot, shaley areas. It was here that the observation was first made that the plants do not commence growth until late July or early August. In addition, these comprise part of the best genus community for Spiranthes in New England. Five species of Spiranthes are present as well as several other species of orchids. Additional orchid taxa noted: Spiranthes cernua, Spiranthes lacera var. lacera, Spiranthes ochroleuca, Spiranthes romanzoffiana, Spiranthes ×borealis; Epipactis helleborine, Liparis loeselii, Malaxis unifolia, Gymnadeniopsis clavellata var. ophioglossoides, Platanthera lacera, Platanthera dilatata, Platanthera huronensis, Platanthera aquilonis, Platanthera psycodes, Platanthera ×andrewsii. Flowering plants of Spiranthes casei observed: 1989 - 41; 1990 - 23 (many mowed spikes); 1991 - 47; 1992 - 67; 1993 - 74; 1994 - 32; 1995 - 28; 1996 - 44. Site 18. Creampoke ii Stewartstown Hollow - Roy residence This site, which is only a short distance from site 17, has afforded the best opportunity for observations in a controlled mowing environment. It consists of a roadside site that continues up the driveway to a grassy hillside. The landowners have been more than cooperative in adjusting their mowing schedules for the areas with the Spiranthes and it has resulted in considerably more flowering plants in 1996. No other Spiranthes species are present at the lower site; however, at the site at the top of the hill Spiranthes cernua, Spiranthes ochroleuca, Spiranthes romanzoffiana and Spiranthes ×borealis are found. Additional orchid taxa noted: Spiranthes cernua, Spiranthes ochroleuca, Spiranthes romanzoffiana; Platanthera dilatata, Platanthera aquilonis, Platanthera lacera, Platanthera psycodes, Platanthera ×andrewsii,
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Flowering plants of Spiranthes casei var. casei observed: 1993 - 4; 1994 - 6; 1995 - 8 at lower site/ 4 at new site at top of hill; 1996 - 13 at lower site/ 11 at top of hill. VERMONT All of the Vermont sites are consistently small populations, rarely subject to mowing and often the victim of succession. Several new sites were found in 1996 during a systematic search of roadsides in areas with suitable habitat. Orleans County. Site 19. Sutton Located on the east side of US Route 5A in the vicinity of a major spring on rocky outcrops and talus slopes of Mt. Pisgah along Lake Willoughby. This entire slope along Lake Willoughby is a rich botanical area. It has always been a good spot for abundant Spiranthes romanzoffiana, Spiranthes cernua and a few Spiranthes ochroleuca. With the exception of 1996, a few flowering plants of Spiranthes casei var. casei could always be found. There appeared to be no disturbance of the area in which Spiranthes casei var. casei formerly occurred and repeated trips failed to disclose any plants that year. Additional orchid taxa noted: Spiranthes cernua, Spiranthes ochroleuca, Spiranthes romanzoffiana, Spiranthes ×borealis; Liparis loeselii, Platanthera dilatata, Platanthera huronensis, Platanthera aquilonis. Flowering plants of Spiranthes casei var. casei observed: 1992 - 2; 1993 - 4; 1994 - 3; 1995 - 5; 1996 - 0. Site 20. Brownington i This was the first recent site for Spiranthes casei var. casei to be found in Vermont (1990). It is on the west side of US Route 5A, 1.4 km north of Vermont Route 58 on an east-facing slope along an access road to a pasture. The slope is densely overgrown with a variety of grasses and Solidago spp. Plants of Spiranthes casei var. casei tend to be short, few-flowered, and restricted to very small open areas within the grassy tufts. The plants here have been the most difficult to see due to the competition from the grasses. Additional orchid taxa noted: Spiranthes ochroleuca. Flowering plants of Spiranthes casei var. casei observed: 1990 - 3; 1991- 4; 1992-1993 - 0; 1994 - 3; 1995 - 0; 1996 - 4. Site 21. Brownington ii A single flowering plant of Spiranthes casei var. casei was discovered in 1996 just south of site 20 on US 5A on a dense, grassy, east-facing slope. No additional species of orchids were noted. Flowering plants of Spiranthes casei var. casei observed: 1996 - 1 Site 22. Brownington iii This site, on the northeast side of Vermont Route 58 on a southwest facing slope 0.75 km west of Vermont Route 5A, is one of the two largest extant sites in Vermont (site 25 being the other). With the exception of 1995, it has consistently produced flowering plants each year. No additional species of orchids were noted. Flowering plants of Spiranthes casei var. casei observed: 1990 - 9; 1991 - 11; 1992 - 4; 1993 - 15; 1994 - 4; 1995 - 0; 1996 - 11. Site 23. Charleston Careful searching of US Route 5A north of the Lake Willoughby region yielded only one
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additional site for Spiranthes casei var. casei at pole 25/35. This low, shaley ridge, with Thuja occidentalis sprouts, is on the west side of the road, 3 km south of the junction with Vermont Route 105. Plants observed were unusually large and vigorous. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1996 - 6. Site 24. Newport This is the northernmost observed site in Vermont, less than 1.5 km south of the Canadian border. The site is on US Route 5, 0.7 km north of the intersection of East Main Street & Sias Avenue at pole 46/39. The plants are growing in a large roadcut of shaley and sandy outwash. Additonal taxa of orchids noted: Spiranthes cernua, Spiranthes ochroleuca. (It is interesting to note that Spiranthes ochroleuca is not known from Quebec but grows here on a very short distance from the border). Flowering plants of Spiranthes casei var. casei observed: 1996 - 8. Site 25. Barton i Located on US Route 5 on a west-facing slope 6.5 km south of Vermont Route 16 in Barton, this site is particularly interesting because it is one of the least disturbed sites in Vermont. The steep, grassy slope is composed of shale with small exposed pockets. It is within these pockets that Spiranthes casei var. casei is found. In 1996 it was the largest site in Vermont. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed : 1990 - 6; 1991- 3; 1992 - 7; 1993 - 4; 1994 - 0; 1995 - 7; 1996 - 12. Site 26. Barton ii A small site located just north of the railroad overpass on a shaded roadside and continuing along to a sunny, exposed, shaley driveway banking. Two Spiranthes casei var. casei and one Spiranthes ×borealis on the roadside; on the driveway curve six Spiranthes casei var. casei, three Spiranthes ×borealis; seven Spiranthes ochroleuca. Few plants of Spiranthes casei var. casei, but excellent representation of the nothospecies Spiranthes ×borealis. Additional orchid taxa noted: Spiranthes ochroleuca, Spiranthes ×borealis. Flowering plants of Spiranthes casei var. casei observed: 1996 - 8. Site 27. Greensboro This site was one of the most gratifying of all those found during the searches made in 1996. In May of 1995 suitable roadside habitat for Spiranthes casei var. casei was observed along Vermont Route 16 in this area. During a return to search these roadsides in 1996, a steep, shaley slope was noted just north of Stannard and south of the Flagg Pond access area. Although plants could not be seen from the vehicle a careful search on foot revealed several plants of Spiranthes casei var. casei near the top of the east facing slope within young Thuja occidentalis. Extensive searching of similar slopes along the highway failed to reveal any more plants of Spiranthes casei var. casei. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1996 - 7. Caledonia County. Site 28. Lyndon US Interstate Route 91 just south of the Lyndon/Lyndonville interchange contains numerous, thin-soil, rocky outcrops within the center strip. In 1990 plants were observed with field-glasses
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that appeared to be Spiranthes casei var. casei. In subsequent years no other plants were seen and in 1996 extensive searches were made of the roadsides in this area. This may be another example of repeated mowing decimating the population. No additional orchid taxa were noted. Flowering plants of Spiranthes casei var. casei observed: 1990 - 12+ plants observed from vehicle; 1991-1996 - 0. Essex County. Site 29. Lower Waterford One of the most extensive stretches of shale roadsides can be found along US Interstate I-93 at the New Hampshire/Vermont border. After careful and repeated searching of several miles of these roadsides, most of which proved to be too wet for Spiranthes casei var. casei, two plants were found on the southbound side of the Interstate, 3.75 km northwest from the New Hampshire state line. Both plants were small, but well flowered. Additional orchid taxa noted: Liparis loeselii. Flowering plants of Spiranthes casei var. casei observed: 1996 - 2. COMMENTS ON SPIRANTHES CASEI VAR. CASEI IN NEW YORK AND PENNSYLVANIA NEW YORK In the course of studying Spiranthes casei var. casei, additional sites were sought out and documented in both New York State and Pennsylvania. These two areas differ greatly and represent two important components of the Location of Spiranthes casei var. casei. The habitat of the populations seen in northern New York State are more typical of those found in the upper Great Lakes region where the body of the populations lie (Case 1987). In the vicinity of Speculator, New York,—both to the southeast and southwest—extensive Spiranthes populations occur. Although some populations were smaller, the majority observed were in excess of fifty flowering plants of Spiranthes casei var. casei and there were many more plants of Spiranthes cernua and Spiranthes ochroleuca. The very distinct, single-ranked hybrid of Spiranthes cernua and Spiranthes ochroleuca was also seen (Catling 1989; Sheviak 1991) (Figure 21). In this geographic area Spiranthes casei var. casei was most abundant in open, dry, sandy grasslands and less frequent on disturbed roadsides. In those sites where both Spiranthes casei var. casei and Spiranthes ochroleuca were sympatric, a few individuals of Spiranthes ×borealis could be found. Although road widening has decimated a large area that previously supported extensive populations of Spiranthes casei var. casei (as well as several other species of Spiranthes) the relatively undisturbed grasslands southwest of Speculator on New York Route 10 have large, healthy populations. Roadsides were carefully searched from the intersection of the New York Thruway (US Interstate 90) and New York Route 10 north to Speculator and southeast toward Lake George. The Spiranthes casei var. casei populations were confined to an area of within 35 kilometers of Speculator. Two superior genus communities of Spiranthes were noted. Each contained from three to five species of Spiranthes, one nothospecies and often several other orchids including Liparis loeselii, Gymnadeniopsis clavellata, Platanthera huronensis, Platanthera psycodes, Platanthera lacera, and Pogonia ophioglossoides. The habitats here varied from streambanks and seeps to the aforementioned open sandy plains and were very different from those where Spiranthes casei var.
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casei was observed in northern New England, where the species was confined to calcareous, shaley barrens and slopes. The reports of specimens of Spiranthes casei var. casei from southern New York State and nearby northern Pennsylvania present well removed disjunct populations. The rediscovery of these plants from one of the Pennsylvania sites in 1996 gave an opportunity to observe this disjunct population. PENNSYLVANIA In mid-September of 1996 an excursion was made to northern Pennsylvania, carefully following directions to three small colonies of Spiranthes casei var. casei that had been found the previous week. After a short search the plants were found on thin-soil roadsides adjacent to a railroad bed. The colonies varied from one to twenty-two plants. They were typical Spiranthes casei var. casei and exhibited no characters that differed from plants elsewhere within the main range of the species. Spiranthes cernua was present, in a wet ditch, nearby to the largest population and a few plants of Spiranthes ochroleuca were found as well as a single Spiranthes ×borealis. After examining the three sites to which directions previously were given, an abandoned railroad crossing and siding near Bingham, Pennsylvania was approached and additional plants of Spiranthes casei var. casei, Spiranthes cernua, Spiranthes ochroleuca and Spiranthes ×borealis were found. What was so very interesting at these sites was their proximity to the rotting railroad ties and the presence of adventive Thuja occidentalis. In northern New England Spiranthes casei is always accompanied by Thuja. The railroad ties in this locality appear to have been hewn from a gymnosperm rather than an angiosperm and the presence of the Thuja suggest that the ties were made from this wood and that the Thuja was present as the result of adventitious burls. At any rate, this disjunct population was associated with a most unusual adventive species common to the plants further north. At one of the sites, the flowering plants of Spiranthes casei var. casei were growing directly out of the rotting ties. Recent documented reports of Spiranthes casei var. casei from southwestern Wisconsin (Hapeman 1996; Weber 1996) have shown plants that have flowers much larger and whiter than are typically seen from Michigan and Ontario eastward. Examination of these plants may reveal gene introgression from the diploid Spiranthes magnicamporum or from a prairie race of Spiranthes cernua.

SPIRANTHES CASEI VAR. NOVAESCOTIAE IN NOVA SCOTIA

Spiranthes casei var. novaescotiae represents an isolated, distinctive population. When Catling & Cruise (1974) originally described the species they noted that it was partially apomictic (op cit., p. 535). This reference referred to plants collected in Nova Scotia (Catling, pers. com.). The following is based upon the original description of Catling (1981). Additional notes are intended to supplement the description. Spiranthes casei Catling and Cruise var. novaescotiae Catling. Figure 2 Canadian Journal of Botany 59(7): 1253-70. 1981. Terrestrial herbs, 0.7-2.8 mm in diameter, tall, basal leaves two to seven, 60-130 mm long, 7-12 mm wide; stem erect, the basal sheaths leaf-bearing, upwardly becoming bract-like; inflorescence terminal, densely flowered, 17-81 mm long; bracts lanceolate to ovate-lanceolate, acute to acuminate, often elongate, white towards the tip, 8-14 mm long; flowers creamy to
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yellowish-green, glandular-pubescent outside; dorsal sepals decidedly concave inwardly, ligulateoblong, slightly constricted above the base on both sides, acute, 5.0-7.7 mm long, 2.7-3.7 mm wide; lateral sepals slightly oblique, oblong-lanceolate, acute to subacuminate, 5.2-7.6 mm long, 2.0-3.0 mm wide; petals ovate-elliptic from a cuneate base, obtuse, distal margin erosedenticulate, 3.9-6.3 mm long, 1.7-2.3 mm wide; lip ovate-elliptic, with a basal claw 0.4-0.8 mm long, the expanded region near the claw obtusely ovate with curved and triangular, fleshy calli, the entire lip 4.1-6.5 mm long, 3.2-4.3 mm wide; column 2.8-3.6 mm long; anther 1.9-2.7 mm long, 0.8 -1.4 mm wide. TYPE: CANADA: NOVA SCOTIA: Shelburne Co.: Hope's Lot Barrens, north of the hamlet of Clyde River (4341'N, 6530'W), 26 August 1975, P. M. Catling & S. M. McKay (holotype, TRT 212892; isotypes, ACAD, AMES, CAN, MICH, TRT 212891). REPRESENTATIVE SPECIMENS: CANADA: NOVA SCOTIA: Shelburne Co.: Hope's Lot Barrens, Clyde River, 7 September 1921, M. L. Fernald & B. Long 23727,23728 (AMES 85418, PH 589277, PH 907414); a few kilometers north of Jordan Falls (4350'N, 6514'W), 25 August 1975, P. M. Catling & S. M. McKay (TRT); 2-3 km west of Jordan Falls (4348.5'N, 6515'W), 25 August 1975, P. M. Catling & S. M. McKay (ACAD, AMES, TRT 212895); 1.6 km W of Jordan Falls (4348.5'N, 6515'W), late August 1978, P. M. Catling, F. H. Catling, & P. F. Maycock (AMES, CAN, MICH, TRT 212890, WIS). Yarmouth Co.: Pubnico, 6 September 1920, M. L. Fernald, B. Long, & D. H. Linder 20918 (AMES 21117, PH 907427). Belleville, 5 September 1921, M. L. Fernald & B. Long 23724, 23726. Also known from Queens County. Spiranthes casei in Nova Scotia is represented by several records of a very distinct autogamous taxon, Spiranthes casei var. novaescotiae Catling, and two records of plants assigned to Spiranthes casei var. casei. Catling (1981) described in great detail the taxonomy of the autumnflowering Spiranthes from southern Nova Scotia and proposed the variety novaescotiae, citing several historical and extant collections at the time of his work. As with several cited specimens of Spiranthes casei var. casei, the specimens that were to become Spiranthes casei var. novaescotiae were previously identified as Spiranthes ochroleuca (Catling 1981; Catling and Cruise 1974). He gave detailed information as to localities of all of his collections. In an attempt to document the current status of Spiranthes casei in Nova Scotia, the province was visited in early September of 1995. Essentially following the pattern of Catling‘s collections, the foray started at Digby on the west side, and continued around to Yarmouth and north to Liverpool and then west back to Digby. All of Catling‘s sites were searched as well as those of Fernald from nearly seventy years earlier (Catling 1981). All three autumn-flowering Spiranthes were found: Spiranthes cernua, Spiranthes ochroleuca, and Spiranthes casei. In addition, all suitable habitat for Spiranthes that was seen during the drive was field-checked. Spiranthes cernua is an apogamous compilospecies found throughout the eastern United States and adjacent Canada, whereas Spiranthes ochroleuca and Spiranthes casei var. casei are both sexual through all of their range excluding Nova Scotia (Catling 1981; Sheviak 1982, 1991). In Nova Scotia both Spiranthes cernua and Spiranthes ochroleuca are apogamous. This greatly simplifies identification due to the fact that introgression and hybridization are not currently present within these three species of autumn-flowering Spiranthes in Nova Scotia.
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Proceeding south from Digby toward Yarmouth, several excellent sites with numerous flowering plants of both Spiranthes cernua and Spiranthes ochroleuca were encountered. But the first plants of Spiranthes casei were not found until habitat along the eastern shoreline northeast of Pubnico was examined. This conformed with Catling‘s previous experiences. It was very late in the day, nearly dark, and the few plants found were not at all typical of either Spiranthes casei var. novaescotiae or Spiranthes casei var. casei. Early the next morning, returning to Nova Scotia Route 103 northeast of Pubnico, a detailed search for additional Spiranthes casei var. novaescotiae sites was begun. The exploration continued along the route, eventually coming to the site seen the previous evening. It was determined to stop at all likely habitat and record which species of Spiranthes, if any, were found. At the first dry roadside banking an abundance of typical Spiranthes casei var. novaescotiae, Spiranthes ochroleuca, and in the wetter ditches at the base of the bank, Spiranthes cernua, were found. This site was still some distance from the plants seen the previous evening. Driving northward toward the previously discovered site, scattered plants of typical Spiranthes casei var. novaescotiae, along with Spiranthes ochroleuca and occasionally Spiranthes cernua, were found. The plants first observed were relocated and then it was realized that perhaps what had been seen the previous evening were Spiranthes casei var. casei. Catling (1981) states that the one collection he referred to Spiranthes casei var. casei was not wholly typical of var. casei but more closely intermediate in morphology between var. casei (away from Nova Scotia) and var. novaescotiae. To prepare for this possibility a carefully collected inflorescence of Spiranthes casei var. casei from Stewartstown, New Hampshire was brought for comparison. Inflorescences of nearby typical var. novaescotiae were collected from the present site. The resulting comparison showed exactly what Catling had described and it was easily seen that the large plants that were found here were obviously intermediate between var. casei and var. novaescotiae. The most interesting fact was that this site was far removed from Catling‘s only site for the intermediate, which was near Brooklyn, Queens County and the current one was to the northeast of Pubnico, Yarmouth County. To date these are the only two records of this intermediate taxon. Although Catling assigned his specimens of the intermediate plants to var. casei this can create more confusion in sorting out what actually occurs in Nova Scotia. Both var. casei and var. novaescotiae are listed for the province. In view of this intermediate plant, typical Spiranthes casei var. casei does not appear to be present in Nova Scotia. Like many taxa in the genus, Spiranthes cernua, Spiranthes ochroleuca and Spiranthes casei var. novaescotiae all colonize disturbed, often roadside, habitats. The lack of competition derived from periodic mowing, coupled with the fact that the species reaches flowering size from seed in only two to four years, often results in extensive flowering colonies, which, although often shortlived, are none-the-less impressive. Not one of Catling‘s specific sites yielded any Spiranthes casei var. novaescotiae. In the twenty years since Catling‘s field work all of his sites had succumbed to succession and were densely overgrown or had been developed. However, there was no difficulty in finding an abundance of Spiranthes species in other suitable habitat. It was estimated that we observed in excess of ten thousand flowering stems of the three species of Spiranthes in three days of field work. Approximate numbers showed the proportions to be: Spiranthes cernua 40%, Spiranthes casei var. novaescotiae 35% and Spiranthes ochroleuca 25%. Roadside bankings, disturbed barrens, power lines, and borrow pits yielded the greatest number of Spiranthes casei var. novaescotiae. Two sites had in excess of one thousand flowering
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plants. Scattered plants were even observed in front lawns. Elsewhere the genus community concept can be applied to Spiranthes communities. Here, where all three species are non-sexual, that application is less relevant. Mixed communities were frequent and actually made identification somewhat easier because of convenient comparison. Spiranthes casei var. novaescotiae is well established, if not locally abundant, along the east coast of southwestern Nova Scotia. It can be easily determined from Spiranthes cernua and Spiranthes ochroleuca by its smaller, yellowish flowers and its often single-ranked spiral. The single clump of intermediate plants seen were slightly larger, with flowers more open and with fewer leaves. To one who is not familiar with Spiranthes casei var. casei they certainly would key to that taxon. Familiarity with both varieties clearly places the Nova Scotian Spiranthes casei var. casei midway between typical Spiranthes casei var. casei and Spiranthes casei variety novaescotiae. In all probability it represents the remnant of sexual Spiranthes casei var. casei on the peninsula prior to the species adaptation to more xeric sites and lack of pollinators.

Fig. 2. Spiranthes casei var. novaescotiae Pubnico, NS
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GENUS COMMUNITIES The concept of genus communities was set forth in 1983 by Wagner and Wagner to deal with mixed populations of morphologically similar species of Botrychium (Ophioglossaceae), the grape ferns. The authors described mixed populations of species that rarely interbred and individuals that maintained their characteristics that defined the species. Although it is not at all unusual to find several species of orchids growing together, there are few genera that have multiple species in the same section that form large communities. When we find several species of Cypripedium growing in a common site, they are of different sections so the concept of genus communities does not apply. It is not that unusual to find Cypripedium acaule, Cypripedium arietinum, Cypripedium reginae, and Cypripedium parviflorum var. pubescens all growing in a common community; however; they are all members of different sections of the genus Cypripedium and are all morphologically distinct. The genus Platanthera is a better candidate for the concept, although here species from different sections also often grow together; large communities of several species are unusual. In the genus Spiranthes mixed communities of similar species are the norm rather than the exception. There is no sectional division in the strictest sense. Northern New England offers several excellent examples of these genus communities. It is not all unusual to find sympatric roadside sites for Spiranthes cernua, Spiranthes ochroleuca, and Spiranthes romanzoffiana in northern New England. Hybridization is rare and the species remain distinct. The following is perhaps the best example of a Spiranthes genus community that can be cited from the Northeast. It also is an exceptional orchid community (Brown 1996).1 Along a winding road in northern New England, not far from the Canadian border, exists one of the most unusual orchid communities in the Northeast. Along this stretch of harsh, sunny, rocky roadside thirteen species of orchids co-exist. This inhospitable roadside contains woodland edges, shaley barrens and a small, intermittent seep. These habitats create ecological niches that altogether cover about 100 meters along the road. Five species of Spiranthes are to be found here. The most notable is Spiranthes casei var. casei, which is found locally in this part of northern New England. The roadside curve has one of the larger populations. In 1996, sixty-two individuals were seen in bloom. Spiranthes romanzoffiana is present in small numbers and Spiranthes lacera var. lacera has been seen, both flowering in August, and soon to be followed by Spiranthes casei var. casei. By September, both Spiranthes cernua and Spiranthes ochroleuca are in full bloom. Spiranthes ×borealis (Spiranthes casei var. casei × Spiranthes ochroleuca) is occasionally seen. Apart from the Spiranthes, five species and one nothospecies of Platanthera are present. Platanthera aquilonis and Platanthera huronensis share the seep with Gymnadeniopsis clavellata var. ophioglossoides. Scattered plants of Platanthera lacera and Platanthera psycodes are along the roadside in the shadier areas as well as Epipactis helleborine and a very unusual yellow-flowered form of the broad-leaved helleborine, Epipactis helleborine forma luteola P.M. Brown (Brown 1996). Several examples of Platanthera ×andrewsii (Platanthera lacera × Platanthera psycodes) can be found. A few scattered plants of Liparis loeselii bring the list to thirteen species and two nothospecies. A short
1

The following originally appeared as A Most Unusual Orchid Community in the North American Native Orchid Journal 1996. 2(3): 275-80. 228

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distance down the road is a large wetland with an abundance of Platanthera psycodes and Platanthera aquilonis. Across the road from the primary site, on the cooler and shadier side of the road, are several dozen Epipactis helleborine in a wide variety of colors from green through cranberry as well as several additional Platanthera aquilonis. I have long been a champion of roadside botany and this particular site is one of the best examples I can present. It was found during a systematic search of the area for new colonies of Spiranthes casei var. casei. As I visited the site more frequently the other species began to present themselves. Several miles south on this same road are large colonies of Spiranthes ochroleuca and Spiranthes cernua, the very distinctive hybrids between them, as well as scattered sites of Spiranthes casei var. casei (Catling 1983; Sheviak 1982, 1991). Malaxis unifolia is also found elsewhere on this same roadway. OBSERVATIONS ON THE PHENOLOGY OF SPIRANTHES CASEI VAR. CASEI. In pursuing studies of Spiranthes casei var. casei in northern New England several known populations were revisited in the spring of 1992 to chart the progress the plants made throughout the growing season. After careful searches no plants could be located at sites that had supported several flowering plants in previous seasons. By late August flowering plants were present. The following year, 1993, these, and similar sites, were again revisited in June and no vegetative plants were found to be present. That fall I marked flowering plants to attempt to trace their growth the following season. Again, in the spring of 1994, no plants were present. By early August leaf growth was just commencing and along with it the flower spikes were evident. In comparing this with other known species of Spiranthes it was noted that all other species of Spiranthes in New England had a very different vegetative/flowering cycle. In most species the leaves were evident very early in the spring, and the genus is one of the first herbs to commence growth. Depending on the individual species of Spiranthes, the leaves made their full growth and either senesced prior to flowering, coincident with flowering or well after flowering. But in all instances, the other species of Spiranthes initiated growth in the spring. At the several genus community sites for Spiranthes casei var. casei in northern New England these other species of Spiranthes were in evidence from late May onward. No plants of tagged Spiranthes casei var. case were evident until late July or early August. This may account for it not being possible to find plants of Spiranthes casei var. casei earlier in the season. As leaf growth commenced so did the flowering spike. Leaf and spike growth proceeded at a proportional rate until the first flowers opened and leaf growth stopped. Discussions with colleagues in other parts of the range of Spiranthes casei var. casei confirmed a similar situation, although not with the exact same observation (Shriver, Weber, Hapeman, pers. comm.). In each instance information indicated that populations in New York, Pennsylvania and Wisconsin no longer were extant when fieldchecked for plants in early summer. By September flowering plants were evident and the field workers assumed they just missed the plants earlier. In reality the plants were not visible, but very much alive. The season of 1995 was one of extreme drought in northern New England. Many species did not do well and flowering was poor among several of the orchids. Spiranthes casei var. casei, however, flowered very well in most sites. At that time, the question arose as to whether this retarded development was simply a survival function to cope with the drought. The season of 1996 was just the opposite. Exceptional rains provided much more moisture than usual. The
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Spiranthes casei var. casei plants behaved in exactly the same manner. The plants were a bit larger and lusher, but the timing was the same as in previous years. Observations were made on individual plants in both xeric and mesic habitats in four populations in two states. All plants commenced growth at the same time. An observation made by Scott Weber (pers. comm.), of Baraboo, Wisconsin, who raises many plants of Spiranthes casei var. casei under cultivation, that they often flower in the greenhouse or beds in late June rather than in the fall supports the observation that the leaf growth and the flowering spike commence simultaneously. His plants under cultivation are cared for in a very different manner than those growing under natural conditions: this has altered their life cycle by inducing growth earlier in the spring, but the flower spike is also initiated earlier and the plants grow at the same rate as those in the wild. HYBRIDS AND NOTHOSPECIES Although several species of Spiranthes form the occasional rare hybrid only three nothospecies have been described. These three occur with more frequency where the parents are sympatric and often have led to confusion in determining plants in the field. Spiranthes ×intermedia Ames Fig. 4 (Ames 1903; Catling 1978) is the result of hybridization between Spiranthes lacera var. gracilis and Spiranthes vernalis (Figure 30). The parents often occur together throughout their range but only in the Northeastern United States do they flower at the same time. The nothospecies was originally described by Ames although there has been much confusion concerning the epithet intermedia. Catling writes in great detail to resolve this confusion and clearly demonstrates that Spiranthes ×intermedia Ames is a nothospecies and not a synonym for Spiranthes casei Catling & Cruise. (Catling, 1978). In Luer (1975) Spiranthes casei is addressed as Spiranthes intermedia. Spiranthes ×simpsonii Catling & Sheviak (Spiranthes romanzoffiana × Spiranthes lacera var. lacera) was described by Simpson and Catling (1978) and was without a formal designation until Catling and Sheviak (1993) completed the work. In this case, the parents also are sympatric throughout much of the northeastern portion of their range, but the nothospecies is a rare occurrence. In much of the existing range in New England Spiranthes lacera var. lacera has finished flowering well before Spiranthes romanzoffiana commences flowering. Only in extreme northern New England do flowering times overlap. Spiranthes ×borealis P.M. Brown Fig. 3 (1995) was first noted as robust Spiranthes casei (or in some situations as small-flowered Spiranthes ochroleuca) in some of the original sightings of Spiranthes casei in northern New England. Although published in 1995 without significant supporting data, the nothospecies was not unknown to others. Both Catling and Sheviak (pers. comm.) had indicated that they had seen such a plant in mixed colonies of Spiranthes ochroleuca and Spiranthes casei var. casei. The following data clearly demonstrate that these plants are the resulting hybrids of the two putative parents and, unlike Spiranthes ×intermedia and Spiranthes ×simpsonii, occur in nearly every sympatric situation in northern New England as well as in New York State and northern Pennsylvania. In both Spiranthes ×intermedia and Spiranthes ×simpsonii, the Spiranthes lacera (in variety) parent imparts the green throat to the nothospecies. In the case of Spiranthes ×borealis, both parents are creamy-white in color and therefore the floral shape, stance and overall morphology are the best indicators. The reddish, gland-tipped hairs characteristic of Spiranthes casei var. casei,
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and lacking on Spiranthes ochroleuca, are imparted to the nothospecies. However, they are not so intense in coloring as in Spiranthes casei var. casei. Additional reports of putative hybrids involving Spiranthes casei var. casei include Spiranthes casei var. casei × Spiranthes romanzoffiana from northern New Hampshire (two plants, same population at site 18, Creampoke ii) and reports of Spiranthes casei var. casei × Spiranthes magnicamporum from southwestern Wisconsin (Hapeman, pers. comm.). Although Spiranthes casei var. casei is sympatric with Spiranthes lacera var. lacera in several northern New England populations, no apparent hybrids have been observed. Because all populations of Spiranthes casei var. novaescotiae are apomictic, as well as all populations of Spiranthes ochroleuca and Spiranthes cernua in Nova Scotia, the likelihood of hybrids there is slight. Both Spiranthes lacera var. lacera and Spiranthes romanzoffiana in Nova Scotia are sexual, and reports of Spiranthes ×simpsonii are beginning to surface, but both of those species usually finish flowering well before Spiranthes casei var. novaescotiae commences.

Fig 3. Spiranthes ×borealis, West Milan, NH

Fig. 4. Spiranthes ×intermedia, South Sandwich, MA

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Literature Cited: Ames, O. 1903. Natural hybrids in Spiranthes and Habenaria. Rhodora 5: 261-64, plate 47. ____1905. Contribution towards a monograph of the American species of Spiranthes. Orchidaceae, fascicle 1: 113-56. ____1921. Notes on New England orchids, - Spiranthes. Rhodora 23: 3-85, plates 127-29. ____1924. An Enumeration of the Orchids of the United States and Canada. American Orchid Society, Boston, Massachusetts. Brackley, F.E. 1985. The orchids of New Hampshire. Rhodora 87: 1-117. Brown , P.M. 1993. A Field and Study Guide to the Orchids of New England and New York. Orchis Press, Jamaica Plain, Massachusetts. ____1995. New taxa - Spiranthes ×borealis. North American Native Orchid Journal 1(4): 290-91. ____1997. Wild Orchids of the Northeastern United States. Cornell University Press, Ithaca, N.Y. Case, F. W. Jr. 1964. Orchids of the Western Great Lakes Region. Cranbrook Institute of Science, Bulletin 48. Bloomfield Hills, Michigan. ____. 1986. Orchids of the Western Great Lakes Region. Cranbrook Institute of Science, Bulletin 48. Bloomfield Hills, Michigan. 251 pp. revised edition. Case, F. W., Jr. and P. M. Catling. 1983. The genus Spiranthes in Michigan. Michigan Botanist 22: 79-92. Catling, P. M. 1978. Taxonomic notes on Spiranthes casei Catling and Cruise and Spiranthes ×intermedia Ames. Rhodora 80(823): 377-89. Catling , P. M. 1981. Taxomomy of autumn-flowering Spiranthes species of southern Nova Scotia. Canadian Journal of Botany 59(7): 1253-70. Catling, P. M. 1989. Biology of North American representatives of the subfamily Spiranthoideae. in Proceedings of the North American Native Orchid Terrestrial Conference, Brandywine Conservancy, Chadds Ford, Pennsylvania. Catling, P. M. and J. E. Cruise. 1974. Spiranthes casei, a new species from northeastern North America. Rhodora 76(808): 526-36. Catling, P. M. and C. J. Sheviak. 1993. Taxonomic notes on some North American orchids. Lindleyana 8(2): 78-79. Chapman, W. K. 1997. Orchids of the Northeast. Syracuse University Press. Syracuse, N.Y. Correll, D. S. 1950. Native Orchids of North America north of Mexico. Chronica Botanica Co., Waltham, Mass. Donley, J. F. 1963. The Orchids of Nova Scotia. J. F. Donley, Mill Village, Nova Scotia. Hapeman, J. 1996. Orchids of Wisconsin. http:/www.wisc.edu/botany/orchids.html Homoya, M. 1994. Orchids of Indiana. University of Indiana Press, Indianapolis, Indiana. Luer, C. A.. 1975. The Native Orchids of the United States and Canada excluding Florida. New York Botanical Garden. Bronx, New York Mousley, H. 1924. Further notes on the orchids of Hatley, Stanstead Co., Quebec, 1923. Canadian Field-Naturalist. 38: 62-63. _____1941. Two orchids new to the Province of Quebec and the Dominion of Canada, Spiranthes vernalis Engelman & Gray and Spiranthes cernua (L.) L.C. Richard var. odorata (Nuttall) Correll. Canadian Field-Naturalist. 55: 79-80 + 2 plates. _____1942. The spring ladies‘-tresses, (Spiranthes vernalis Engelmann & Gray) in Canada.
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Canadian Field-Naturalist. 56: 1-2 + 2 plates. Munden, C. 1995. Orchidizing in Nova Scotia. North American Native Orchid Journal 1(3): 171-94. Pease, A. S. 1964. Flora of Northern New England. New England Botanical Club. Cambridge, Mass. Seymour, F. C. 1969. The Flora of New England. Charles E. Tuttle Co., Rutland, Vermont. Sheviak, C. J. 1982. Biosystematic study of the Spiranthes cernua complex. New York State Museum Bulletin No. 448. Albany, New York. ______. 1991. Morphological variation in the compilospecies Spiranthes cernua (L.) L.C. Rich.: Ecologically limited effects of gene flow. Lindleyana 6(4): 228-34. Sheviak, C. J. and P.M. Catling. 1980. The identity and status of Spiranthes ochroleuca (Rydberg) Rydberg. Rhodora 83(832): 526-62. Simpson, R. C. and P.M. Catling. 1978. Spiranthes lacera var. lacera × S. romanzoffiana, a new natural hybrid orchid from Ontario. Canadian Field-Naturalist 92(4): 350-58. Voss, E.G. 1972. Michigan Flora. - Part 1. Gymnosperms and Monocots. Cranbrook Institute of Science & University of Michigan, Bloomfield Hills, Mich. and Ann Arbor, Mich. Wagner, W.H. Jr. and F.S. Wagner. 1983. Genus communities as a systematic tool in the study of new world Botrychium (Ophioglossaceae). Taxon 32(1): 51-63. Weber, S. 1996. Spiranthes cernua and Spiranthes casei on a farm in Wisconsin. North American Native Orchid Journal 2(1): 57-63. Whiting, R.E. and P.M. Catling. 1986. Orchids of Ontario. CanaColl Foundation, Ottawa, Ontario. Acknowledgements: The many years spent preparing this paper involved information from the following persons: Chuck Sheviak, Paul Catling, Scott Shriver, Jeff Hapeman, Scott Weber, Shirley Curtis & Sally Puth, Debbie Benjamin, and numerous private landowners; Stan Folsom aided in all of the field work.

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Empiricist: WHATEVER GOT YOU INTERESTED IN ORCHIDS?

WHATEVER GOT YOU INTERESTED IN ORCHIDS?
The Slow Empiricist
From my position of advancing years I am still interested in a lot of things and curious about a great number of things. As I ponder about the possible topics for my Slow Empiricist column I began thinking about what sparked my interest in orchids. This led me to wonder what some of the readers of my column experienced as they became interested in orchids. To help you focus on your own story I have documented some of the things that shaped my interest in the native orchids. My love of plants seems to have been inborn in me. I probably got the genes, if there are such genes, from my maternal grandfather who was a Down East farmer and lobsterman. He grew a big vegetable garden that helped feed not only his own family but others in the neighborhood. His vegetable garden always had room for flowers such as marigolds, bachelor buttons, cosmos, and incredibly beautiful sweet peas in an endless bloom of dazzling colors. Unfortunately for me I lived in a city suburb of Boston, Massachusetts although we had room for a flower border and our yard had two old pear trees and a young elm and maple. Most of my gardening was tending the perennials such as the iris, phlox, hollyhocks, and daylilies. I did have a small Victory Garden during the Second World War. I was only in elementary school but I grew all the easy vegetables like radishes, carrots, lettuce, and Swiss chard, but I did not have peas, corn, or tomatoes. After the war and rationing ended we were able to visit our summer home in Maine as well as my grandparents Down East. I was a preteen but I began to enjoy the natural world around me and I encountered my first native orchids although I did not know them as orchids. The woods around my cottage had pink lady‘s-slippers and a vernal stream bank had yellow lady‘s-slippers. I don‘t remember picking them because I was used to flowers like daisies and brown-eyed Susan's which made nice bouquet material, which was most propitious for the lady‘s-slippers continued existence. They are abundant today here in Acton, Maine. By the time I was an adult I grew the garden variety of plants and shrubs that I liked but orchids weren‘t among them. Living in the Northeast was not conducive to my growing exotics and I was basically ignorant of the orchid family. I had gone to art school, received my teaching credentials, and got established in my career. I had even visited the Peabody Museum at Harvard University and seen the magnificent glass flowers, but was still untutored about orchids or plant families. For the most part orchids were exotic hothouse flowers used in corsages. It wasn‘t until I met PMB and started attending some of his field trips did I learn about our native orchids. By then I was middle aged and open to new experiences. The idea of seeing all the native orchids was an interesting challenge and has provided me with many years of travel and excursions to explore and find these plants. As I march toward my 75th birthday, I have seen
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most of the native orchids in North America north of Mexico. I say most, because new species are being found and described even as I write! I have been in all 50 states and most Canadian provinces and territories finding orchids all along the way from the high mountains outside of Tucson in August, to the open forests outside of Anchorage in June to the streams at the base of Mt. Albert in the Gaspé in eastern Canada to most anywhere orchids grow. Now that I live in Florida, I can see orchids every month of the year from slender coralroots in January to grass-pinks in March to the fringed orchids in the summer to the ladies‘tresses practically all year long! My interest grew then out of an initial love of nature and gardening and only needed the right combination of conditions to blossom into a full blown activity that has enriched my life. I think it would be very interesting to hear what set of circumstances caused you to be passionate about wild orchids. Do you have a story to share? Your Slow Empiricist

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