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NORTH AMERICAN NATIVE ORCHID JOURNAL
(ISSN 1084-7332) published quarterly in March June September December by the NORTH AMERICAN NATIVE ORCHID ALLIANCE, Inc. a group dedicated to the conservation and promotion of our native orchids Editor: Paul Martin Brown Assistant Editor: Nathaniel E. Conard Editorial Consultants: Philip E. Keenan Stan Folsom Production Assistant: Nancy A. Webb The Journal welcomes articles, of any length, of both a scientific and general interest nature relating to the orchids of North America. Scientific articles should conform to guidelines such as those in Lindleyana or Rhodora. General interest articles and notes may be more informal. Authors may include line drawings, and/or black and white photographs. Color inserts may be arranged. Please send all inquiries or material for publication to the Editor at PO Box 772121, Ocala, FL 34477-2121 (mid June - August: PO Box 759, Acton, ME 04001-0759). 1999 Membership in the North American Native Orchid Alliance, which includes a subscription to the Journal, is $26 per year for United States addresses, $29US in Canada and $32US other foreign countries. Payment should be sent to Nancy A. Webb, 84 Etna St. Brighton, MA 02135-2830 USA.

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NORTH AMERICAN NATIVE ORCHID JOURNAL
Volume 3 Number 2 CONTENTS NOTES FROM THE EDITOR
138

June 1997

PREPUBLICATION ANNOUNCEMENT THE GENUS CYPRIPEDIUM 140 WHY DO PLANTS CHANGE NAMES?
P. M. Brown 143

CYPRIPEDIUM PARVIFLORUM VAR. PUBESCENS IN THE SOUTHWEST
R. A. Coleman

150 SAPROPHYTIC ORCHIDS OF DALLAS
V. S. Engel

157 FIELD TRIP ETIQUETTE
The Slow Empiricist

168 HISTOLOGICAL AND DEVELOPMENTAL STUDIES OF THE SHOWY LADY'S-SLIPPER CYPRIPEDIUM REGINAE
K. Sokolski, A. Dovholuk, L. Dovholuk,K. Lavigne, K. Ganey & P. Faletra

198 THE GENUS CYRTOPODIUM IN FLORIDA
R. L. Hammer

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ORCHID SEARCH - MANITOBA 1996
L. Heshka

187 ALL TRESSED UP, BUT NO PLACE TO GO or Local Ladies’-tresses Orchids of Ohio
T. A. Sampliner

215 BOOK REVIEWS Wild Orchids of the Northeastern United States Orchids of the Northeast Orchids of the Ottawa District, Ontario Orchids of the Bruce and Grey Counties, Ontario Hidden Orchids 238 LOOKING FORWARD September 1997 248 Color Plates 1. Cypripedium parviflorum var. pubescens 2. Hexalectris nitida, Hexalectris warnockii 3. Cyrtopodium paranense 4. Platanthera praeclara

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 3, number 2, pages 138-250; issued June 16, 1997. Copyright 1997 North American Native Orchid Alliance, Inc. COVER: Dactylorhiza aristata var. kodiakensis by Stan Folsom

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NOTES FROM THE EDITOR
As we continue into our third year, this issue presents a wide variety of articles from all over North America. Our first article from the Canadian prairie provinces is included as well as our first from Texas. Many of you have traveled widely in North America and I urge you to write something for a future issue. The March issue devoted to the genus Cypripedium was very well received and I welcome all the new NANOA members who joined as a result of purchasing single issues. We are continuing to reprint that issue to fill the demand of single orders. Because of the time demands of filling and printing individual issues this will be the only single issue available. Full sets of back issues for both 1995 and 1996 are still available and will continue to be so. Contact Nancy Webb for more information on back issues. Ron Coleman‟s excellent article in Orchids, March 1997, concerning our first conference in Pittsburgh was very well received and garnered several new members. We thank the American Orchid Society for its generous coverage of the Alliance and our activities. Do not forget our second conference to be held in Arizona this summer. There is still time to register.

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An information flyer was included with the March issue. It is time to prepare for a 1998 calendar. The subject will be the fringed orchises of North America. Our new color process will ensure a better quality of reproduction and this time ONLY vertical, 8X10” color, glossy, prints will be considered. You may either send the finished prints or slides. If I am aware of slides that individuals have, I may be contacting you for consideration. The calendar will be finished by September, so all slides or prints to be considered must reach me before August 1, 1997. I trust you will continue to have a rewarding and productive field season and look forward to your many reports. Please remember the summer address is PO Box 759, Acton, Maine, 04001-0759 and the phone/fax is 207/636-3719. Paul Martin Brown editor naorchid@aol.com

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PREPUBLICATION ANNOUNCEMENT
Coming in August 1997 from Timber Press

THE GENUS CYPRIPEDIUM
by Phillip Cribb In this monograph, distinguished author Phillip Cribb has considered the history, biology, evolution, conservation, cultivation, and classification of the temperate slipper orchids of the genus Cypripedium. Orchidists, particularly those interested in slipper orchids and terrestrial orchids, as well as other botanists and woodland and alpine gardeners will appreciate this richly illustrated and highly authoritative volume. Cypripediums, popularly called lady's-slippers or moccasin flowers, are the showiest and most sought after hardy orchids, collected and grown by orchid and alpine plant enthusiasts alike. They have been used medicinally in North America and the Far East for centuries, having been grown as far back as 2,500 years ago during the time of Confucius in China. Interest in these plants revived in the nineteenth century when plants from Siberia, North America, and the Himalayas were sent to Europe, where the Industrial Revolution gave people the time and

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resources to build glasshouses in which to grow exotic plants. Slipper orchids have fascinated horticulturists and botanists alike, being set apart from other orchids most notably by a prominent slipper-shaped lip. Their pollination biology was first studied by Charles Darwin. The major source of these plants has been from the wild; thus many of the more accessible populations of many species have declined through overcollection. In Europe and North America, other factors leading to extinction include habitat destruction, particularly logging and agricultural improvement, and drainage of natural habitats. The only British species, also that country's rarest plant, has been reduced in its range to a single known locality and, tragically, to a single plant. This clump has survived alone in its protected site in northern England for more than fifty years. Yet the lady'sslipper was common enough in the area in the nineteenth century to be collected for table decorations in a local hostelry and to be dug up to adorn gardens in the region. Most of the striking members of the genus are natives of North America, growing well with ferns in shaded, damp, acid soils rich in organic matter. Although several species are thought to be difficult to grow, the author clearly identifies plants suitable for beginners and, for all species, provides formulas for various mixes used successfully by growers of

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Cypripediums. No other book on these important orchids exists at present. Phillip Cribb is curator of the Orchid Herbarium at the Royal Botanic Gardens, Kew. He is the author of several books on orchids, including The Genus Paphiopedilum and, with co-author Ian Butterfield, The Genus Pleione, both in this series. He has traveled widely in connection with his work and has studied the genus Cypripedium in the wild on several trips to China in the past twenty years. Approx. 300 pp, 26 full-page color paintings, 98 color photographs, 51 black-and-white illustrations, and 22 maps, 6 x 9", hardcover, ISBN 0-88192-403-2 Price not set, Publication date: August 1997. Contact: Deborah Garman, Publicity Manager. Available from Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527. Telephone Nos. (800) 327-5680, (503) 227-2878, Fax (503) 227-3070. http://www.timber-press.com/ and email at publicity@timber-press.com

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Brown: WHY DO PLANTS CHANGE NAMES?

WHY DO PLANTS CHANGE NAMES? Paul Martin Brown
The change of plant names after years of general usage often confuses plant enthusiasts. Why plants change names is, perhaps, the most frequently asked question I receive from members and students alike. The basic answer is quite simple; the reasons behind it are not. Plants change names for one of two reasons: the first is that the currently used name is either incorrect or invalid—each species has only one validly published name; the second is that recent research has shown that the specific species is in need of a new name. How? When? Why? Where?????? When a specific plant is to be identified, it has both a genus and a species name given to it. These names are in Latin and are the same for all botanists worldwide. This two-part name is known as a binomial. Binomial nomenclature is the naming of a plant with two names: a genus name (the group of similar plants) and a specific epithet (the name which sets off the species). The name of the plant consists of both the genus and specific epithet. For brevity the

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word „species‟ is often used in lieu of „specific epithet.‟ Prior to the work of Linnaeus in the mid-1700s, plants (and animals, etc.) had a variety of Latin names in various combinations, but not in a standard format. Pre-Linnaean names were often long and cumbersome, sometimes consisting of 10-15 descriptive terms. In an effort for uniform classification, Linnaeus, in his Species Plantarum of 1753, devised a system of assigning a genus name and a specific epithet to each distinct individual or group of individuals. The Latin, or scientific, name of the plant consists of a genus and a specific epithet followed by the name of the author, the first person to validly describe the plant. This is what we know as plant taxonomy: the classification of plants, including their naming, or nomenclature. Please keep in mind that we are discussing plants here, specifically orchids. All of the following guidelines apply to all plants, as opposed to other organisms (birds, mammals, etc.) which may have somewhat different rules. For the next 150 or more years it was not unusual for a single species to receive more than one name, as different botanists each thought they were the first to describe the plant. This was particularly true of variable plants of wide distribution. The name that was used the most became the most widely accepted name for that species. With the formation of the

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International Association of Plant Taxonomists in the early 20th century, it became apparent that a set of well-defined rules were necessary to standardize plant names. These eventually became the International Code of Botanical Nomenclature, often referred to simply as the Code. New editions of the code are issued following each international conference. The latest was in 1995. Among many other things, the Code sets forth a series of universally agreed upon guidelines for: (a) naming new species, transferring names from one genus to another, or creating subspecies, varieties and forms; and (b) more importantly, what constitutes a valid publication of a new name or combination. The Code is very simple: the earliest validly published name takes precedence. Sounds easy. Very often it takes years of researching the literature to determine what is the earliest validly published name. Rules have been formulated that must be met to insure a plant name‟s validity. Since 1935, to be validly published, a species (or subspecies, variety or form) must have a Latin diagnosis (a brief statement in Latin stating the significant differences from similar taxa) and have a lengthier description in the language of the author; it must cite a type, which is the specimen from which the original description is made. The type specimen must be deposited in a permanent herbarium.

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In addition, it must be effectively published. That is, it must be published in either a journal or major work on the genus that is a permanent work (not a newsletter) and that is distributed and available to the professional botanical community from which the species is described. It is unfortunate that several good species were not validly published and therefore their names cannot be used (see NANOJ 1(3):254-266 concerning the taxonomy of the Kentucky lady’sslipper, Cypripedium kentuckiense). If a Latin diagnosis is lacking, that omission invalidates the publication; if the author‟s name is missing, that omission invalidates the publication; or, if no type is designated, that omission invalidates the name. If a Latin name is used in the literature prior to publication and not identified as in ed. ( a work that is in progress but not yet published), it too can invalidate that name for future use. But back to names changing. Prior to 1900, new species did not always have a full Latin description or diagnosis, which complicates choosing the earliest, valid name for the plant. Careful research of the literature is the primary culprit for changing the name of a popular plant to a less well known or even obscure species name. According to the nomenclature rules, if a name is correct (i.e. a prior publication that has been validly published) it must be used. But it can often be awkward. An example of this can be seen in the genus Malaxis, the adder’s mouths. Malaxis macrostachya (Lexara) Kunze is correctly known as M. soulei L.O.

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Williams, because William‟s publication predates Kunze‟s. Upon occasion when a species is deemed to be too broad and two or more species are the result of the research, the original name can only be applied to the correctly identified type specimen. The use of the more familiar name may or may not be employed for the most widespread and/or familiar resulting species. Malaxis ehrenbergii (Reichb. f.) Kunze is widely distributed in Mexico but the plants of northern Mexico and adjacent Arizona, New Mexico, and west Texas were determined to be a new species, Malaxis wendtii Salazar (see NANOJ 2(1):65-66. Name changes at the generic level have the same rules but here we have two guiding forces. One is, of course, the correct or valid name. The other force is taxonomic opinion. This is the result of research that some taxonomists clearly feel demonstrates that the genus is in need of division and more than one genus is the result. Sometimes, it simply utilizes prior genera and the species are grouped according to their technical characters; alternatively, a new genus is described to accommodate those species that do not fall into existing genera. Which genera to use and which species belong to each genus are very often matters of opinion. Each taxonomist tries very hard to state his or her case and some will agree and some will not. This is a nuisance, especially when a well-known and

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familiar species has been changed. An effort has been made to reduce this by a list of conserved genera, i.e. genus names which cannot be changed even if an earlier name is found. There is however, no provision for conserving specific epithets. In the North American native orchids, the two groups in which we see this segregation happening the most, are the fringed and rein orchises, Habenaria and the ladies’-tresses, Spiranthes. For the past half century the genus Habenaria has been a large, allinclusive one that accommodated many species. Recent works, especially like that of Carlyle Luer, have demonstrated that several of the original genera that were merged into Habenaria are more appropriately retained as separate genera. Hence, we now see Platanthera, Coeloglossum, Pseudorchis and Piperia being used. These are not necessarily new names, but simply the earlier names for smaller groups of species (see NANOJ 2(1):28-28 for details). Another interesting example is the genus Orchis. Two species were always well known from North America: showy orchis, Orchis spectabilis and small round-leaved orchis, Orchis rotundifolia. Again, relatively recent work in the mid-20th century demonstrated that neither of these species actually had the critical characters that the Eurasian species of Orchis possessed. In this case the earlier name of Galearis accommodated Orchis spectabilis but the creation of a new genus, Amerorchis, was needed for

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O. rotundifolia. The plants do not change, only their names. Paul Martin Brown

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Coleman: CYPRIPEDIUM PARVIFLORUM VAR. PUBSCENS IN THE SOUTHWEST

CYPRIPEDIUM PARVIFLORUM VAR. PUBESCENS IN THE SOUTHWEST Ronald A. Coleman
By the end of the summer of 1995, my struggle to find the large yellow lady‟s-slipper, Cypripedium parviflorum var. pubescens, in Arizona had proceeded in vain for three years, and showed no signs of abating. In early 1996, I decided to expand my horizons somewhat and look for it in the neighboring state of New Mexico. The large yellow lady‟s-slipper has historically been more numerous in New Mexico than in Arizona, and might, therefore, prove easier to find. My motivation was that knowledge gained by studying the plant in habitat there might yield some clues on where to look for it in Arizona. Several weeks of research turned up multiple documented locations within six to eight hours of driving time from my home. Tom Todsen, an orchid researcher who lives in New Mexico, agreed to go with me. On 25 May we met for lunch in a town part way to our destination, and caravaned the rest of the way. We made a brief stop en route to explore one of Tom‟s favorite orchid

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places, and found spotted coralroot, Corallorhiza maculata, and a Schiedeelia species in bloom. We also found Wister’s coralroot, Corallorhiza wisteriana, in fruit. That evening we camped by a small stream and planned the route for the following morning. Tom had visited this lady‟s-slipper colony many years earlier, and had a pretty good idea of how to get there. The orchids were in a box canyon, about six miles from the road. We could get there either by going cross country along the ridges, or by following the stream bed. We decided to go in via the ridges, and come out along the stream. Not too far from camp, an informal hunter‟s trail headed into the back country, and we started following it. The first half mile was up hill, but then the trail leveled off in fairly open piñon pine and juniper forest. The walking at first was easy, but after a couple of hours, the trail became faint, and finally disappeared altogether at the base of a large hill. We had noticed a canyon coming in from the left for the last mile or so, and we headed toward it around the side of the hill. The easy walking was over, though, because the hillside was covered with football-sized loose rocks interspersed with several species of cacti. Eventually, we saw in the distance a section of canyon wall that stood out because it was green and lush in sharp contrast to the rest of the mostly brown terrain; we assumed that this was our destination. The last two miles were to take much longer than the first four.

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Between us and the green wall were arms of the canyon that were too steep to traverse, and several times we were forced to back-track up the hill looking for safe passage. Finally, we came to a sheer drop of over 100 feet, but just beyond lay the green section of canyon wall. Off to the side we saw a wooded slope leading down into the canyon. By hanging onto trees and bushes we managed to work our way to the bottom, and then waded and rock-hopped upstream to where we hoped to find the orchids. The green patch, toward which we had painfully worked our way, was an enormous seep about 40 yards wide that reached about 200 yards up the canyon wall. At the stream bed, sections of the seep ended in straight stream-cut bank, while other sections were collapsed piles of mud and muddy boulders. Water was dripping off the entire length of the seep. Ten or twelve feet up, the slope was not nearly as steep, and the seep was covered with grasses and sedges. We scanned the seep from the bottom, and had almost decided we were either in the wrong place, or too early in the season, when I noticed a speck of yellow out of the comer of my eye. Blooming in the grasses only a few inches from the edge of the cliff was a Cypripedium parviflorum var. pubescens. It took several minutes of wet, muddy climbing, but we worked our way up to the orchid. First Tom

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pushed me, than I pulled him, and eventually we reached some soggy but stable ground. The entire seepage area was wet to the touch, and our feet sank in a little at each step. The orchid we had spotted from below was so close to the edge of an area of recent collapse that it seemed certain to fall into the stream during the next major storm. Scattered around the seep were ten additional large yellow lady’s-slippers; most of these were a safer distance from the cliff. Seven of the plants were either in bloom or in bud. The blooming plants all held single flowers, and were between 15 cm (7 in.) and 30 cm (12 in.) tall. The pouches were a bright yellow, and the staminodes were yellow with red dots. The sepals and petals had a base color of green, nearly totally covered with dark brown stripelike markings that turned to scattered dots near the column. Pines and firs lined both sides of the seep, but the orchids were in full sun for portions of the day. The surrounding grasses were nearly as tall as the lady‟s-slippers, and it was very difficult to see the ones that were not in bloom. In an even wetter part of the seep were hundreds of Platanthera in spike. Tom speculated they were Thurber’s bog orchis, P. limosa, because he had seen it elsewhere in the area. We spent some time photographing the orchids and exploring the seep, but then realized it was getting late. The trip back to camp along the stream bed was as difficult as the way in along the ridges. The canyon

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bottom was littered with a jumble of boulders, and we had to cross the stream many times. It would have been impassable in spots had the water level been higher. We did not find any other orchids along the stream, although the habitat looked as if it should support stream orchis, Epipactis gigantea. The trip to and from the seep was long and rough, but thanks to Tom Todsen, I achieved my objective of learning a little more about the habitat of yellow lady’s-slippers in the Southwest. Shortly afterwards, I finally found Cypripedium parviflorum var. pubescens blooming in Arizona. They were growing in grassy, damp soil by a small stream, slightly more shaded but very similar to the habitat in New Mexico. I wish that I could claim the discovery in Arizona was made easier by the study in New Mexico, but it was really just a matter of luck (and perhaps that should be another story).
Ronald A. Coleman

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Engel: SAPHROPHYTIC ORCHIDS OF DALLAS

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Engel: SAPHROPHYTIC ORCHIDS OF DALLAS

SAPROPHYTIC ORCHIDS OF DALLAS Victor S. Engel
In the summer of 1981, I was wandering around the escarpment of southwest Dallas, Texas, in search of something photogenic to put on the last half of a roll of film so I could send it off to be developed. My wanderings brought me to a grove of junipers and oaks. After looking around for a while, I noticed something that looked at first glance like an orchid. I did not know there were any orchids in the Dallas area, so I was quite surprised. On closer inspection, it was clear that I was looking at the stem of an orchid plant laden with seed capsules. There were no leaves or open flowers on the plant. I looked around for others but found none except a smaller plant growing right beside the first one. When I got home, I looked through one of my orchid books and concluded that what I had found was probably a species of coralroot, Corallorhiza. In August of 1982, I went back to the same place to see if the plant had come up again. Unfortunately, it had not. So, I looked around the immediate area for other plants. It was not long before

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I found about a dozen plants full of flower buds and developing ovaries. Only one plant had an open flower, and that flower was open only partially. I looked through several orchid books to try to identify the orchid, but I could not find any matches. Any plant that looked even remotely like the right species had a range that did not include the Dallas area. The following year, I started sooner in my search for the plants in order to catch them in an earlier stage of development. However, instead of finding immature inflorescences of the same species, I found another species in full bloom. I sent a specimen to the Orchid Identification Center at the Marie Selby Botanical Gardens in Sarasota, Florida, and the plant was identified as Wister’s coralroot, Corallorhiza wisteriana, a fairly widespread species whose range includes the Dallas area. The plants I was originally looking for did not come up that year at all. It wasn‟t until July of 1986 that they came up again. This time, I dissected one of the flowers to try to make a proper identification. I noticed that instead of having four pollinia, as Corallorhiza does, it had eight. By that time, I was somewhat familiar with the crested coralroots, the genus Hexalectris, so I concluded that the plant was a species of that genus. The size and shape of the flowers when spread out flat indicated that the plant was the shining crested coralroot, Hexalectris nitida. However, the range was wrong.

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Hexalectris nitida had been reported only in Panther Hill in the Glass Mountains of Brewster County, Texas, and in the Sierra Mojada in the state of Coahuila, Mexico. There were some other problems. In Carlyle A. Luer‟s The Native Orchids of the United States and Canada excluding Florida, H. nitida was described as having open flowers with sepals and petals curving backward toward the stem. The plants in Dallas very rarely had open flowers. Another difference was that according to Luer, there were five ridges on the tip of the lip‟s mid-lobe. The plants in Dallas had five ridges on the lip, but the ridges were not at the tip. Because of the differences, I sent a specimen to the O.I.C. for identification. Soon I got a reply confirming that the plant was an autogamous form of Hexalectris nitida. In Dallas, Hexalectris nitida seems to prefer living in the leaf litter of a mixed juniper and oak forest. Although I have seen some plants in oak litter, the vast majority prefer a bed of decaying juniper needles below which is limestone bedrock. Although all the plants seen were growing near the escarpment, they did not grow past the edge. Presumably, this is because moisture cannot accumulate very well on the sloped surfaces of the escarpment. Also, the soil is not as thick on the slanted surfaces. Corallorhiza wisteriana grows in the same forest as Hexalectris nitida. However, C. wisteriana

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shining crested coralroot Hexalectris nitida

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prefers forest dominated by oaks, whereas H. nitida prefers junipers. Corallorhiza wisteriana also grows in poorer, sandy soil that is populated by grasses. Plants growing in these areas were always smaller than the ones growing in oak litter. Corallorhiza wisteriana also seems more tolerant of sunlight. Plants frequently are seen in clearings in the forest. Another saprophytic species previously unknown in this area is Texas purple-spike, Hexalectris warnockii. When I found the original plant of H. nitida in 1981, I told Dale Williams, an orchid grower who lived near the location of the orchids. I asked him if he had ever seen any native orchids in the area, and he said that he had not. My discovery piqued his curiosity, and after he saw the plant that I had found, he kept his eye out for orchids on a neighboring property that was downwind from the plant that was blooming. In the summer of 1986, he was rewarded not only with many plants of H. nitida but also with a new, showy species, H. warnockii. This species was known to have occurred in parts of central Texas as well as the Big Bend area in Brewster County. Thus, Williams‟ discovery constitutes an extension of the previous range. Another saprophytic species reported in the area in 1986 was the crested coralroot, Hexalectris spicata. They were seen at the Fort Worth Nature Center in Fort Worth, Texas. Dale Williams and I have also seen them on the property of the Dallas Nature

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Texas purple-spike Hexalectris warnockii

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Center (previously known as Greenhills Environmental Center). In the Dallas area, H. spicata blooms in May and June, whereas H. nitida blooms in late June and July. During our studies of the genus Hexalectris in Texas, Dale Williams and I found confusing plants that resembled Hexalectris spicata but were not that species, nor could they be placed with any other currently recognized taxon. Among their distinctive features were relatively small auto-pollinating flowers that rarely opened fully, their columns lacked a rostellum and the keels of the lip were relatively low. Consistently for 10 years, plants of this distinctive taxon have appeared in the Dallas area, but in 1992 I found similar plants in the Austin area of Texas. These plants were eventually recognized as the Arizona crested coralroot, Hexalectris spicata var. arizonica Catling & Engel (with a range from central Texas to southeastern Arizona and adjacent Mexico). At the Dallas site, Hexalectris spicata var. arizonica occurs within an area of oak-juniper woodland that is occupied by both H. nitida and H. spicata var. spicata, whereas at the Austin site it is accompanied by H. nitida. Hexalectris spicata var. arizonica flowers in June and July, later than H. spicata var. spicata which flowers primarily in May (occasionally to early June) in eastern Texas.

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crested coralroot Hexalectris spicata

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Arizona crested coralroot Hexalectris spicata var. arizonica

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Most of the plants seen in the Dallas area and all of those seen so far in the Austin area have closed flowers that never open, but some plants with open flowers have been found at the Dallas site. The closed flowers tend to have more truncate perianth parts than the open flowers. Athough it may be more widespread than is currently known, Hexalectris spicata var. arizonica is nevertheless, based on current information, a geographically restricted taxon. Fortunately, it is protected with other native flora in some of the sites where it occurs.

Editor’s note: This article is based upon two very different earlier works by Victor Engel in the AOS Bulletin 56(8):831835 and by Paul M. Catling and Victor Engel in Lindleyana 8(3):119-126. They both appeared in a different format and have extracts merged here. The previously published material is used with the permission of the American Orchid Society. The latter article is the original publication of Hexalectris spicata var. arizonica and goes into great detail conerning the morphology and potential origins of the variety. The reader is

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urged to consult that reference for full details concerning the variety. VICTOR S. ENGEL, 3305 Yellowpine, Austin, Texas 78757

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FIELD TRIP ETIQUETTE The Slow Empiricist
This is not an easy article to write, but as the spring season has nearly ended and orchid enthusiasts have begun to embark on another series of explorations into the field it is an article whose time has come. Let me begin by stating the obvious—that if everyone treated their fellows with the same courtesies that they would like to receive, then this article would be a waste of time. However, that does not seem to always be the case. If you think the previous statement about courtesy smacks of Biblical overtones you are right. Unfortunately, in our hectic modern world little credence is given to the “golden rule.” You only need to get in your automobile and drive a little slower than the speed limit to find out that human nature is not as tolerant as it perhaps should be. This article has a twofold purpose: first, to make the field trip participants aware of some of their less than admirable behaviors (which may help them become more sensitive to others); and second, to encourage the others to find gentle ways to foster true concern for each other when sharing orchid expeditions.

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If you are alone when you are in the field in your orchid exploration you can have pretty much of a free rein as to how much time you take with the plants and you will not be spoiling someone else‟s pleasure. That is unless you roll on the plants or crush some under foot in your quest to get the perfect shot for your slide collection. It is also common sense that you don‟t disturb the environment so much that you endanger the plants, such as by weeding out the detritus just so you can get an unimpeded view of your subject (some orchids need protective cover). Nor is it such a good idea to leave some kind of noticeable marker to call attention to the species lest some less scrupulous person happen upon the scene and decide to collect the plants for his/her garden or flower vase. When you are with others, however, the rules change. Most people are very considerate of their companions and try to share the site with their neighbors. If you recognize yourself in any of the following situations I would suggest that you might be considered a tad selfish. Do you always have to be the first one to take photographs? Do you always push to the front and hog the best specimens? Are you the first to complain about anything, be it weather, insects, plant material, others‟ behavior or whatever? Are you such an inveterate perfectionist that you have to have at least an hour or more to achieve a satisfactory result? It is to these few that I suggest the following behavioral changes. It would be to everyone‟s benefit

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if the less generous field trip participant were able to recognize his or her bad traits and work to modify them. First, when you get to the site of one of the specialties you have been dying to photograph or commune with ever since you were bitten by the orchid bug, hang back! Let someone else have first dibs on the plants! While you are impatiently waiting for your turn you might try exploring for other specimens. You might be rewarded for your efforts by finding even nicer ones than the original ones you were so eager to get at. If you have really become a sharing, caring companion you will share your discovery right away, even to the point of letting someone else have first go at them. By abandoning thoughtless me-first behavior you may find the rewards are greater than you realize. I urge you to persist. Second, if you are a complainer, then I suggest you thank your lucky stars or whatever you like to give thanks to that you have been able to get out and arrive at the state of being you are now experiencing. There are plenty of fellow enthusiasts who through no fault of their own can‟t be there. They have grown too elderly to make the trip. They have failing eyesight, a case of influenza, or even such mundane problems as job constraints on their time in the field or some other such problem such as an unsympathetic spouse who begrudges their time spent on field trips. Looking at

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your irritations in this way may help you be a better companion to your fellow sufferers for I acknowledge that every field trip doesn‟t always result in wonderful experiences. The weather may be miserable or the insects intolerable or the habitat daunting, but a good attitude will help the situation rather than make it worse. If you go prepared for the unexpected, you will have a better time of it. I always carry a little kit of things that will make my stay a little more pleasurable and help me survive the more nasty elements of nature. In my kit I carry some extra food (in case I wander off the beaten path and believe me I have; or the expedition lasts longer than my stomach can take). I pack a small flask of water in case the weather is too hot for me. I also take a non-aerosol can of bug repellent for I know the best orchids have guardian insects who love my tender flesh. I also tuck in some tissues in case I have a problem that needs wiping or swabbing. If I know the terrain is going to be rough, I carry my walking stick to help me up and over obstacles. I also cover myself with layers of clothing. If it is too cold my layers help protect me. If it gets increasingly warmer, I can divest myself of my layers, draping them about my waist or slinging them over my arm. I always wear a hat. It keeps low branches from tangling my hair and bugs have less of an area to attack. Another item I find indispensable is a small pair of field glasses. They help me locate plants on hillsides and off into the distance without having to

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trudge onto the actual site when I am exploring. There have been many times when I have given silent thanks for having remembered my binoculars. It should go without saying that the avid photographer should not stint on the amount of film or equipment he needs to really do the job. When you are miles from anywhere, running out of film is a real calamity for the photographer. If all my paraphernalia seems a little too much, you can tailor your survival kit to your specific needs. To the inveterate perfectionist, I suggest that you wait to be last with the plants so that you can spend all the time you need to study. One good example of the perfectionist being aware of his fellows is this: when he became aware others were waiting, the photographer allowed others to photograph while he was setting up or in between takes of his. If this situation discombobulates you, find a specimen of your own to work with. If that doesn‟t appeal to you, consider coming back on the morrow to conduct your business. Perhaps you should not participate in group outings if you can‟t speed up your photography time or make some kind of allowance for the others in your party. It should not be for your sole benefit and reward. Group expeditions should be a celebration of the group‟s combined experiences that have been shared. That means accepting the levels of expertise of your fellow members, their physical abilities (some of us

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aren‟t as swift of foot as others or have such sharp eyes as others). It also means being a cheerful companion even if it sounds Pollyanna-ish to look for the bright side in less than perfect situations. It means being helpful and considerate of your fellow celebrants. With attitudes like that you will have pleasant memories of your outing even if it was less than perfect or not up to your standards. You don‟t need to sacrifice your standards. You just don‟t have to make everyone else unhappy because you didn‟t achieve one hundred per cent of your goals. I hope that these messages have reached you in a gentle manner and not offended anyone who suspects that he/she might be seen as a self-centered individual. If you have been guilty of such behavior in the past, you might want to take a harder look at the way you come across. You may find that a few slight changes in your actions might reward you with more camaraderie and a happier experience in the field. If you are one of the patient ones who has been victimized by someone who has been inconsiderate while in your presence I urge you to speak up and assert your rights in a courteous manner—if you can. This might show the errant one good etiquette in the field. If not, you may have to resort to something more drastic as one lady did who had put up with a thousand complaints from a fellow member for over a week. The member had complained about everything.

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It was too hot! It was too cold or too windy! There weren‟t enough plants or there were too many plants! When she complained on the final day that during the entire trip she had not taken one photograph worth her time and money, her companion turned to her and said, “Then you should learn to take better photographs!” Need I say more?
The Slow Empiricist

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A MONOGRAPH OF CORALLORHIZA (ORCHICACEAE)
by John Freudenstein Harvard Papers in Botany no. 10 April 1997. ISSN 1043-4534. 51 pp. $27.00 + $2.00 shipping. For those of us fascinated by the saprophytic genus Corallorhiza it is a pleasure to seen John Freudenstein work come together in a publication that treats the entire genus in greatest detail. Whereas the majority of species are native to North America it is of great interest to many NANOA members. Each species is treated in full detail with line drawings of all the floral parts, distribution maps and extensive lists of representative specimens. For those reader‟s who have been following John‟s work through numerous publications and his dissertation (Cornell 1992) over the past 10 years, this publication pulls all of that work, and more, together in a single highly usable document. (PMB) Copies may be ordered from Carolyn Hesterberg, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138.

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THE GENUS CYRTOPODIUM IN FLORIDA Roger L. Hammer
There is but a single species of Cyrtopodium native to the United States, and that species is Cyrtopodium punctatum (L.) Lindl., colloquially known as the cowhorn or cigar orchid, in allusion to the long, cylindrical pseudobulbs. In the United States, C. punctatum has been recorded on southern, mainland Florida in Broward, Collier, Dade, Lee, Martin, Monroe and Palm Beach counties where it typically inhabits coastal mangrove-buttonwood forests, cypress swamps and mixed hardwood swamps and forests. It can be found growing epiphytically on a variety of trees, especially cypress (Taxodium distichum) and buttonwood (Conocarpus erectus), and is also frequently found perched on the trunks of fallen, dead trees. Although it is commonly found as a terrestrial species in the tropical Americas, it is almost always epiphytic in Florida.

Cyrtopodium punctatum, was first discovered in the United States near Miami, Dade County, Florida in March, 1877 by Abram Paschall Garber, PhD. The species was first described by Carl von Linné (who is better known by his latinized name Carolus Linnaeus)

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in 1759 as Epidendrum punctatum in the publication Systema Alaturae. In 1833 John Lindley transferred it to the genus Cyrtopodium, which he published in his Genera and Species of Orchidaceous Plants. The type locality was not stated but the global range of C. punctatum includes southern Florida, the West Indies, Mexico, Guatemala and Costa Rica southward to Brazil and Paraguay in South America.

When in flower, Cyrtopodium punctatum is a spectacular sight. In late winter each year, new growth and flower spikes appear at the same time from the base of the previous year's leafless pseudobulbs. The inflorescence quickly towers over the leaves and by late March or early April, showy flowers open. Each widely-spreading flower bears greenish-yellow sepals spotted with reddish-brown, bright yellow petals spotted with reddish-brown, and a reddish-brown lip with a yellow base. Each inflorescence may be adorned by thirty or forty flowers and a large specimen will literally display hundreds of blossoms. It has been suggested that the flowers of Cyrtopodium mimic members of the Malpighia Family, Malpighiaceae, and that native bees, during their search for nectar, inadvertently pollinate the orchid flowers. Because of its attractive floral display, the cowhorn orchid has been highly sought by collectors over the years. Photos of wagonloads of cowhorn orchids being brought out of the Everglades just after

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the turn of the present century reveal how common this species once was, but today it is rare and found only in rather remote, protected areas. The historic abundance of Florida's orchids was documented by the famed naturalist Charles Torrey Simpson and his friend, correspondent and renowned botanist John Kunkel Small. In 1916, Simpson and Small went on an orchid collecting trip in southwestern Florida and reported that the orchids were so abundant that their harvest could scarcely be noticed. Cyrtopodium punctatum is the only species mentioned by Dr. Donovan Correll in his publication Native Orchids of North America north of Mexico (Stanford University Press, 1950). Dr. Carlyle Luer, however, in his book The Native Orchids of Florida (The New York Botanical Garden, 1972) reported a yellow-flowered, terrestrial species from Florida which he referred to as Anderson’s cowhorn orchid, Cyrtopodium andersonii (Lambert ex Andrews) R. Brown. Although C. andersonii is cultivated in the Miami area, the species that is naturalized in southern Florida, which Luer photographed, has been determined to be another tropical American yellowflowered, terrestrial species known as Parana cowhorn orchid, Cyrtopodium paranaense Schlecter. The notes of the late field botanist George N. Avery reveal an interesting account of the original reports of this plant in Florida. While attending a Native Plant Workshop meeting in Miami on June 15,

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1971, a woman named Betty McCormack mentioned to Avery that a Cyrtopodium had volunteered inside her screen porch in limestone gravel. Nearly one year later, on May 12, 1972, McCormack brought Avery a portion of her flowering Cyrtopodium, which Avery pressed and deposited in the herbarium at Fairchild Tropical Garden in Miami (Avery #1155). Although McCormack told Avery that she had found a picture of it in Exotica 3 as Cyrtopodium andersonii, Fairchild Tropical Garden's superintendent Stanley Kiem correctly identified it as Cyrtopodium paranaense. Avery pointed out that he had found an article by John Beckner concerning C. andersonii in the Florida Orchidist, the publication of the South Florida Orchid Society (Vol. 7, No. 2, pp. 78-79, 1964), that treated this species as a native, and an earlier issue of the same publication (Vol. 2, No. 4, p. 5, 1959) that featured an article by F. S. Shuttleworth reporting C. andersonii from Florida's west coast. Luer (1972) states that Fred Fuchs, a Miami orchid grower, identified some flowers of an orchid brought to him "from a plant growing terrestrially deep within the Everglades" as Cyrtopodium andersonii. Luer also mentions that "occasionally someone reports finding a specimen [of Cyrtopodium andersonii] in the Everglades, but [it] usually turns out to be a yellow form of Cyrtopodium punctatum." It should be pointed out that there are no verifiable collections of any yellow-flowered Cyrtopodium from the "Everglades," and that the native C. punctatum is the only species

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listed in the extensive botanical inventories for Everglades National Park and the adjacent Big Cypress National Preserve, together encompassing over two million acres. Cyrtopodium paranaense does, however, occur as a naturalized species in at least one natural area—a sandy pineland preserve in the Kendall area of Dade County—where it is firmly established and thriving, even after an intense wildfire in April, 1995. It also readily escapes locally around residential areas wherever it is cultivated in southern Florida and perhaps elsewhere. During several interviews and phone conversations in June and July, 1972, Stanley Kiem gave an interesting historical account of Cyrtopodium paranaense to George Avery and myself. Kiem stated that it was introduced into Florida in ca. 1949-50 by John F. Kasper, who operated an orchid nursery at NW 27 Avenue and NW 15 Street in Miami. Kasper had received a specimen labeled Cyrtopodium paranaense from Sao Paulo, Brazil. Kiem acquired a plant from Kasper in 1951 and cultivated it at his residence at SW 118 Street and SW 94 Avenue in Miami. It escaped as a terrestrial in his yard and as an epiphyte on the prostrate trunks of saw palmetto (Serenoa repens), as well as on a dead pine trunk in a nearby pineland. In September, 1972, Kiem told me that it had spread five miles around his residence in five years. Kasper closed his business and retired at SW 112 Street and SW 92 Avenue in 1954, bringing the Cyrtopodium with him. Three years later, Kiem found plants naturalized at the

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residence of Ed Jordan who lived just a few houses away from Kasper. On May 31, 1976, I sent flowers collected from Jordan's residence to Dr. Kiat W. Tan, then associated with Marie Selby Botanical Gardens in Sarasota. Dr. Tan determined that the flowers belonged to Cyrtopodium paranaense, and made a voucher specimen of the flowers, which he deposited in the Gardens‟ herbarium. A rather painful and disheartening memory of collecting the flowers from Jordan was that of while walking across his yard, I occasionally stepped on crunchy objects beneath my feet. They were mowed-off pseudobulbs of plants that had come up in his lawn! Dr. Gustavo Romero, Research Associate in Orchidology in the Oakes Ames Orchid Herbarium at Harvard University, borrowed specimens from the Fairchild Tropical Garden in May 1991 and reviewed the Cyrtopodium specimens deposited in the herbarium. Dr. Romero attached a note to Avery's 1972 Cyrtopodium paranaense specimen stating that "this material is indistinguishable from Cyrtopodium glutiniferum Raddi," and a label was placed on the sheet to reflect this determination. Cyrtopodium glutiniferum is yet another tropical American, yellow-flowered terrestrial species. In December 1995, I telephoned Dr. Romero in an attempt to clear up this puzzle. Dr. Romero admitted

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to erring when he made the C. glutiniferum determination and that he felt that the Florida plants are correctly C. paranaense. In May 1996, I sent Dr. Romero, by overnight mail, a flowering, wild-collected Cyrtopodium from the Kendall pineland site, as well as flowers of a cultivated, robust, yellow-flowered Cyrtopodium obtained in 1993 from Lynn Bretsnyder of Impact Orchids in Princeton, Florida. Dr. Romero confirmed by telephone that the wild-collected plant was Cyrtopodium paranaense, and that the cultivated specimen was C. andersonii. Dr. Romero has deposited the wild-collected plant in Harvard University's herbarium to further voucher its presence as a wild plant in Florida. Dr. Romero has in preparation an article concerning the genus Cyrtopodium that he plans to submit to the American Orchid Society for inclusion in the publication, Orchids. His work will hopefully put an end to the nomenclature confusion of Florida's naturalized populations of Cyrtopodium, and help identify yellow-flowered members of this genus in the tropical Americas and in cultivation as well. In a letter to me dated 8 April, 1996, Dr. Romero stated "Cyrtopodium paranaense has smaller flowers, and the sepals and petals tend to be roundish. The larger flowers of Cyrtopodium andersonii and Cyrtopodium glutiniferum overlap in size, but the

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labellum is indented and the petals are lanceolate, undulate in C. andersonii versus labellum not indented and petals with a claw, roundish toward the apex in Cyrtopodium glutiniferum." Before finalizing this article for the North American Native Orchid Journal, various reference books were checked to determine the natural range of Cyrtopodium andersonii, and here is the confusing summary: Hortus Third (Staff of the Liberty Hyde Bailey Hortorium, 1976) lists it from Venezuela to Brazil; The Native Orchids of Florida (Luer, 1972) shows a range that includes southern Florida, Cuba, the Lesser Antilles and northeastern South America; The Manual of Cultivated Orchid Species (Bechtel, Cribb & Launert, 1986) has it occurring in the West Indies and tropical S. America; and Garay & Sweet in the Flora of the Lesser Antilles, Orchidaceae(1974) determined that it is endemic to St. Vincent in the Lesser Antilles. To defend their determination, Garay & Sweet state that "Cyrtopodium Andersonii has been confused with Cyrtopodium flavum and Cyrtopodium Engelii by all contemporary authors since Lindley, who united these two different species under Cyrtopodium Andersonii." It is clearly apparent that much work needs to be done on this and other species native to the West Indies and the tropical Americas. In closing, what has been determined to be Cyrtopodium paranaense may likely be in Florida to stay. It has already demonstrated its ability to survive

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fire and periodic, short-duration freezes, it readily produces seed pods, and it is capable of spreading from seed in Florida. All it will probably take is for a storm to move across southern Dade County when seed pods are dehiscing to spread this South American native into other favorable habitats; perhaps even into the Everglades. It may just be a matter of time.
Roger L. Hammer 17360 Avocado Drive Homestead, FL 33030 Gustavo Romero, Orchid Herbarium of Oakes Ames, made helpful suggestions to the manuscript.

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Heshka: ORCHID SEARCH – MANITOBA 1996

ORCHID SEARCH - MANITOBA 1996 Lorne Heshka
I‟ve had an interest in orchids ever since I, as a boy, saw my first yellow lady's-slipper back on the farm in Saskatchewan. At that time there was no doubt in my mind that this was an "exotic" flower and later, when I studied botany at University, I discovered just how unique this family of plants is. This interest, however, remained at an "awareness only" level until an assignment to Thailand with The Food and Agriculture Organization of the United Nations in 1989 and 1990. There I "discovered" tropical orchids and was given, as a gift, several plants. This started me on a new and exciting hobby of growing orchids and I soon found that the slipper orchid species became my favourites. These slipper orchids, the genera Paphiopedilum and Phragmipedium, reminded me of that first orchid I had been fascinated with back on the farm those many years ago. In comparison to the hundreds of orchid species found in tropical countries of the world, the 37 species (and varieties) listed for the Province of Manitoba appear insignificant. The harshness of our winter climate is an important controlling factor, with winter

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temperatures sometimes hovering around -35 degrees Celsius. Prior to 1996, I had observed only 20 of the 37 orchid species that can be found in our province. Although it wasn't a New Year‟s Resolution, early in 1996 I decided to make an effort to find and photograph as many of our native orchids as I possibly could through the year. A couple of weeks of vacation were booked during peak blooming periods, and then the research began. From previous experience, I already knew the location of 20 species. These, however, were the most common and the easiest to find. It was the other 17 that challenged me. Sixty-five dollars‟ worth of text books from the Museum of Man and Nature in Winnipeg ensured that I at least knew what these flowers looked like and the habitat they preferred. Fortunately, I also had several contacts who could give me some specifics as to where and when certain orchids could be found. As I did my literature review, I came to the realization that there were at least three species that I wasn't going to see. These were ones that could be found only in the far north and I had no plans to trek up to Churchill that year. Another species has been found only once before, in the Duck Mountains, and

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has never been seen again. The most I could possibly find, I decided, would be 33 species. The largest number of species of our native orchids occur in the southeast corner of the province, and a large number are found in bogs. As a consequence, my search was planned in the area with the Red River as the western boundary, the Winnipeg River to the north, the United States to the south and Ontario to the east. This area of approximately 25,000 square kilometres has a variety of bogs and other habitats suitable to orchids, and also has a number of Ecological Preserves ensuring minimum human encroachment. The late spring, I quickly found, had delayed flowering and so all of my projected blooming dates were out between one and two weeks. It was, however, an "orchid perfect" summer—ample moisture at the right time, and warm but not scorching weather. Mosquitoes were bad early in the spring, but in the bogs where they are usually the worst, there were few to none of those little pests. I maintained a diary of our excursions and a tally showed that my wife Joan and I covered over 5000 kilometres by car, about 75 km on foot (mostly through bogs), and spent 175 hours searching in every possible habitat. Joan caught the "orchid fever" just as I had and was my constant companion. She became an

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Amerorchis rotundifolia small round-lef orchis

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excellent "bog walker" and it was her sharp eyes that helped us locate some of our rarest finds. On the best day we located 11 different species, the worst day, never fewer than 4 or 5. The weather cooperated with ample periods of overcast, providing excellent light for flower photography. Remarkably, we were never "rained out.” We began our search on June l in the Tall Grass Prairie near Tolstoi located within a few miles of the US border and ended our season there on September 7, coincidentally finding our last species, Great Plains ladies’-tresses, Spiranthes magnicamporum, within a hundred meters of where we found the first, the small white lady’s-slipper, Cypripedium candidum. The "highs" of this project were unquestionably the discovery of several new species (new to us, that is) and meeting some super people along the way. Finding not only one, but several, very rare albino (white-flowered) showy lady's-slippers, Cypripedium reginae forma albolabium, in a localized area had to be the highlight for us—although my wife would argue that point, considering the excitement I displayed when we found for the first time the beautiful and uncommon rose pogonia, Pogonia ophioglossoides. The "lows" were finding spade holes where only a week before a colony of the beautiful stemless

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(pink) lady's-slipper, Cypripedium acaule, stood. The "lowest" wad the news that a political decision had been made to put a well into a bog to raise the water level in a nearby resort lake, effectively sounding the death knell for 19 species of orchids living in that bog. We never did make it to the far north for sparrow’s-egg lady’s-slipper, Cypripedium passerinum and the flat-petalled form of the large yellow lady’s-slipper, C. parviflorum var. pubescens (planipetalum form1). We never found the white adder’s-mouth, Malaxis brachypoda, in the Duck Mountains and the three other small indiscrete species, auricled twayblade, Listera auriculata, northern twayblade, L. borealis, and heart-leaved twayblade, L. cordata var. cordata, eluded us. We had to satisfy ourselves with thirty of a possible thirty-seven species. The new friends we made, the fresh air and exercise we enjoyed and best of all, a tremendous set of transparencies to share with others, made this project worthwhile.

1

The forma planipetalum Fernald is an extreme of the the var. pubescens; see Charles Sheviak‟s article on the yellow lady‟s-slippers in the December 1996 issue of the Journal. Ed.

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Platanthera hookeri Hooker’s orchis

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rose pogonia Pogonia ophioglossoides

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Following is a list of the orchids we observed along with the date they were first found blooming in 1996:
Scientific Name Amerorchis rotundifolia Arethusa bulbosa Calopogon tuberosus var. tuberosus Calypso bulbosa var. americana Coeloglossum viride var. virescens Corallorhiza maculata var. occidentalis Corallorhiza striata var. striata Corallorhiza trifida Cypripedium acaule Cypripedium arietinum Cypripedium candidum Cypripedium parviflorum var. makasin Cypripedium parviflorum var. pubescens Cypripedium reginae Cypripedium reginae Common Name small round-leaved orchis dragon's-mouth grass-pink Venus (eastern fairy) slipper long-bracted green orchid western spotted coralroot striped coralroot early coralroot stemless (pink) lady's-slipper ram's-head lady'sslipper small white lady'sslipper northern small yellow lady'sslipper large yellow lady'sslipper showy lady's-slipper showy lady's-slipper First Observed June 20 June 29 July 1 June 10 June 29 July 7 June 20 June 10 June 10 June 10 June 8 June 15

June 15

June 29 July 7

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(white flowered form) Goodyera repens lesser rattlesnake orchis Goodyera tesselata checkered rattlesnake orchis Liparis loeselii Loesel’s twayblade Malaxis unifolia green adder'smouth Platanthera dilatata tall white northern var. dilatata bog orchis Platanthera hookeri Hooker's orchis Platanthera hyperborea tall leafy green var. hyperborea orchid Platanthera lacera ragged fringed orchis Platanthera obtusata blunt-leaf orchis forma foliosa Platanthera orbiculata large round-leaf orchis Platanthera praeclara western prairie fringed orchis Platanthera psycodes small purple fringed orchis Pogonia rose pogonia ophioglossoides var. ophioglossoides Spiranthes lacera var. northern slender lacera ladies'-tresses Spiranthes Great Plains ladies'magnicamporum tresses Spiranthes hooded ladies'romanzoffiana tresses

forma albolabium

July 27 July 20 July 22 July 7 July 7 June 10 June 20 July 28 June 20 July 11 July 6 July 27 July 14

July 28 September 7 July 31

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Postscript - Because of the efforts of a close friend who was unafraid to challenge the bureaucracy, the installation of the well has been delayed until an environmental impact study is completed. We are hopeful that the public outcry, as a result of the media attention given to this issue, will result in the consideration of other environmental-friendly solutions. Lorne Heshka, 1204 DeGraff Place, Winnipeg, Manitoba, R2G 1Y8 CANADA notes on the photos:
The western prairie fringed orchis, Platanthera praeclara, photograph was taken in the Tall Grass Prairie adjacent to the US boundary southeast of Winnipeg. 1996 was an exceptional year for this species and the counts in this managed area for this year exceeded the total for the previous three years together. Over 20,000 plants were counted and this truly is a success story which may be worthy of follow-up. The white-flowered form of the showy lady’s-slipper, Cyripedium reginae forma albolabium, is found in the localized area threatened by the well mentioned in my article. A rough estimate is that approximately 5 to 10% of the population of Cypripedium reginae in this area are the white flowered form. I haven't had the opportunity to fully explore the extent of this colony with respect to this area. The blunt-leaved rein orchis, Platanthera obtusata, photograph demonstrates a characteristic I have observed in this species only once previously. Part way up the flower stalk is a smaller, thinner leaf (forma foliosa). This small group of plants exhibit this characteristic while others in the same location were typical in that they had only the single leaf.

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Sokolski et al.: Cypripedium reginae

HISTOLOGICAL AND DEVELOPMENTAL STUDIES OF THE SHOWY LADY'S-SLIPPER CYPRIPEDIUM REGINAE Katie Sokolski, April Dovholuk, Lauren Dovholuk, Kim Lavigne, Karen Ganey and Peter Faletra
The showy lady's-slipper, Cypripedium reginae, is a terrestrial orchid found across the northern regions of central and eastern North America. In New Hampshire this species commonly grows in marshy areas or fens preferring the more alkaline soils found on the border of Vermont and New Hampshire. Although these plants have never been found in great numbers in New England, owing to both habitat restrictions and loss, they are now critically endangered in much of northeastern North America. Cypripedium species require the help of insects for pollination. Orchids, in general, are rather specific as to the insects which they will accept as pollinators. In some cases orchids can only be fertilized by the

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males of a single species of insects. However, this is not the case in lady's-slippers (Withner, 1974). The physical arrangement of these flowers creates a tight space through which the insects (mostly bees) have to squeeze. A pollinating insect first passes by the stigma, and upon exiting the trap rubs against the anther. The insect that now carries pollen must reenter another lady‟s-slipper to successfully carry out pollination (Dressler, 1981). One study suggests only twenty out of one thousand bumble bees are willing to pass this gauntlet twice. When successfully pollinated, like many orchids, Cypripedium seed pods contain thousands of seeds. Because the seeds do not contain endosperm (Dressler, 1981), they seem to require a more rigorous set of conditions to germinate and develop than seeds with endosperm. One of these conditions is the presence of symbiotic fungi. Although Cypripedium species, such as C. reginae and the large yellow lady’s-slipper, C. parviflorum var. pubescens, are rare, when they do occur, they can often be found in great numbers, concentrated in a small area. The habit of the flower to occur in this colonial fashion seems, in part, owing to their multiplication via rhizomes. The showy lady’s-slipper is often regarded as the most regal of the genus with two flowers (rarely three or four) occurring on threefoot tall plants. Cypripedium reginae takes about 8 years to mature to a flowering state in the wild (Withner, 1974). Therefore, any approach which speeds this

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maturational process may prove to be an important adjunct to restoration attempts. In vitro culture could be of extreme value since it can speed this process considerably. Nevertheless, for in vitro culture to be of practical worth a variety of challenges with regard to efficiency must be met. When considering efficiency, in vitro methods which affect germination percent and speed, along with subsequent development and survival, are central factors. Chu and Mudge (1994)2 obtained acceptable germination rates for C. parviflorum (as C. calceolus) when seeds were surface sterilized in bleach for twelve minutes. Studies on C. reginae (Dovholuk and Faletra, 1996; Dovholuk et al., 1997) showed an increase in percent germination when exposed for up to thirty-six minutes in bleach. Various other approaches have been attempted to maximize efficiency (Faletra et al., 1997; Sokolski and Faletra, 1997). The general goal of this investigation was to devise a method of efficiently producing large numbers of Cypripedium reginae through in vitro culture and micropropagation. Large scale in vitro culture could then supply sufficient numbers of seedlings for a restoration attempt. To this end, two specific approaches were used: 1) Axenic seed culture, 2) Micropropagation of seedling tissue.
2

Chu and Mudge address the plants they used as Cypriepdium calceolus var. pubescens, an older and incorrect name for C. parviflorum var. pubescens. See Sheviak, C.J. 1996. NANOJ 2(4): 319-343 for full taxonomic details. Ed.

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Axenic seed culture could supply seedlings for restoration along with sterile tissue for micropropagation investigations. Studies of the morphology, histology, and development were done in hopes that the normal processes of the seedling development could serve as a guide to successful micropropagation. These investigative approaches were aimed at answering the following questions: Where are the meristematic tissues? What, if any, tissues are determined and what are they destined to become? At what point can determined meristematic tissue be manipulated? What are the areas of the seedlings which can be micropropagated? How can in vitro raised seedlings be transplanted and raised to normal flowering plants? Can this approach help alleviate the decline of an endangered species in the face of habitat loss? Axenic seed culture studies Conditions that affect speed of germination and subsequent development affect the efficiency of the system. With respect to germination, since each pod produces thousands of seeds, seed supply, although important, is not a primary factor with regard to efficiency. With an average plating density of about

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45 seeds per culture (although we usually see over 90% germination) a greater than 10% germination rate should give quite an adequate number of seedlings per culture vessel. Figure 1 shows results from experiments on germination rate and subsequent development in seeds exposed to bleach for increasing amounts of time. With germination beginning as early as 12 days, a steady progress in efficiency was seen in increased rates of germination and speed of development of the seedlings at longer exposure times to bleach. The differences among the groups were significant at p<.05. These and following experiments (data not shown) indicate that exposures between 36 and 60 minutes in bleach give optimal germination. Rates of development reinforce this trend. The decrease in stage three seedlings seen in Fig. 1 is most likely owing to their development into later stages which show increasing numbers with increased exposure to bleach. Interestingly, this and other studies had never tried simply soaking seeds in water as a control, since without any sterilent, cultures would have little chance of growing free of contaminants. This is unfortunate, since as the diluent for bleach, H2O is the appropriate control. Preliminary results employing a sterile distilled water (SDH2O) control show extended time in H2O up to sixty minutes increases germination rate and development, but it is not yet certain whether the water alone is as effective as bleach. Further studies are in progress. An option would be to soak seeds in

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SDH2O prior to, or after surface sterilization. This is also being investigated. An alternative to solid medium is liquid medium. A disadvantage of liquid medium is the reputed need for rotation which requires an expensive orbital mixer. Therefore, it seemed reasonable to investigate the need for rotation. Seeds placed in nonrotating liquid culture were not as quick to germinate and did not have as high a germination percentage as seeds in rotating liquid culture. This may be attributed to more effective oxygenation and elimination of wastes in the rotating cultures in addition to a more even osmotic pressure. These results suggest there is a distinct advantage to rotating liquid media. Our early studies and those of Chu and Mudge (1994) showed liquid media to support higher germination rates than solid medium when seeds were sterilized at 12 minutes in bleach. Since Chu and Mudge used Cypripedium parviflorum it was of interest to perform a similar experiment on this species. Preliminary results indicate that, as with C. reginae, increased time in bleach up to 36 minutes increased the germination rate and rate of development for C. parviflorum. We have also found that the germination and development of Cypripedium reginae in solid medium, for seeds exposed to bleach for more than 36 minutes, is about the same as for seeds in liquid

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medium. The longer surface sterilization seems to level the differences between solid and liquid media. Consequently, although a liquid medium might give slightly better percent germination, if the speed of germination and development is not substantially greater than solid medium there seems little advantage in the liquid; especially considering the need for an orbital shaker. Histological and Developmental Studies Because axenic culture of Cypripedium species has only recently been successful, studies of morphology and development have been somewhat superficial. This is unfortunate since development is a key to understanding any plant. Scanning electron microscopy (SEM) consistently reveals an orifice at only one end of the seed (Fig. 2 and 2A). This hole could possibly be a consequence of the incomplete closure of the micropyle during seed development. The opposite end was probably attached to the placenta and bears no opening. Within the seed coat is contained an embryo of approximately 500 cells enclosed in a coat possibly derived from interior integuments (Fig. 3). Seed germination studies indicate that in order for germination to proceed this coat needs to be penetrated by what is most likely water.

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It was of interest to determine the fate of seedling tissues. Figure 4 shows the various tissues of a seedling approximately ten weeks old. Roots were revealed by the presence of a root cap and unicellular root hairs (Fig. 4A). Shoots were indicated by the presence of stomata (Fig. 4B). Light microscopy (Fig. 6 and 6A) shows that early seedlings are composed of undifferentiated parenchymal cells with no cotyledons which is typical of orchids but atypical of the class Monocotyledonae (Scagel, 1965). The cells of the seedling shown in Figure 6A possess nuclei in a dense arrangement indicative of meristematic tissue in the region we refer to as the apex. It is meristematic tissue that is a logical target for micropropagation. We have monitored this area over a period of months in a number of seedlings and in all cases have seen it develop into a shoot. This has also been supported by SEM which shows this tissue to contain developing stomata. Cross and longitudinal sections of root are shown on Figure 7 and 7A respectively. The presence of central vascular tissue made up of xylem and phloem indicate that this is root tissue. The unilateral positions of starch granules and nuclei probably indicate the horizontal orientation of this root. The presence of large stores of starch also indicate that these roots store substantial amounts of energy. Figure 8 shows a longitudinal section of shoot tissue opposed on either side by a sheath which is believed to be a coleoptile-like

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structure. This reinforces our belief that Cypripedium grows much like grasses, with the exception of slipper orchids having successive roots emerging at nodes and rhizome tissue making up the internodes. This will occur until the plant is stimulated to produce an aerial shoot (see drawing). From these and future studies that localize and explain the development of both meristematic and committed tissues, successful micropropagation will be more probable. Micropropagation Unfortunately, the genus Cypripedium has not yielded to attempts at micropropagation. Hoshi et al. (1994) recently published results of a fairly exhaustive micropropagation attempt, revealing marginal success in the way of limited callus production and no evident protocorms. Our micropropagation attempts with similar approaches to that of Hoshi et al. (i.e., matrix experiments) have not been successful. Histological sections of early seedlings localized meristematic tissue (Fig. 6). This seemed important for success in micropropagation. Unfortunately, when these young seedlings were sectioned, no success in micropropagation occurred. This is possibly owing to the difficulty in cutting such small specimens. Nevertheless, attempts to micropropagate by

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sectioning older seedlings were somewhat successful. As discussed above, scanning electron micrographs were taken to positively localize different committed tissues. Knowing exactly where root and shoot tissue were located helped guide sectioning. When 2-6 month old seedlings were cut approximately in half, with a piece of a shoot and one or more roots on each half, both halves developed into healthy seedlings and eventually matured to the level at which they were ready to be vernalized (Fig. 9). In some cases, young seedlings could be cut in more than three pieces. Older and larger seedlings, being easier to section and possessing more shoots, often yielded more than three sections. Although this approach might technically be considered as micropropagation, it is not the most desirable situation. Ideally, micropropagation generates generous numbers of protocorms grown from tissue randomly cut from a single plant's tissue. We are now attempting some novel approaches to micropropagation and are also working to improve the methods for axenic culture of other endangered orchid species. We also hope to be planting a considerable number of showy lady’s-slippers in their natural habitat this coming year.

We would like to thank Ms. Elaine Faletra for her help with Light microscopy, Dr. Ezequiel R. Rivera for his histological assistance and Dr. Charles Daghlian for his help in SEM. This

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research was funded in part by a grant from the Toyota Motor Co.

Peter Faletra, PhD, 59 Mountain Meadows Rd, Warren, NH 03279
Literature Cited Chu, C. & Mudge, K. 1994. Effects of Prechilling and Liquid Suspension Culture on Seed Germination of the Yellow Lady‟s-slipper Orchid. Lindleyana 9(3): 153-159. Dovholuk, A., Faletra, P. 1996. Development of Cypripedium Species in Plant Tissue Culture. Bulletin of American Association for the Advancement of Sciences, Annual Meeting 1996. Dressler, R.L. 1981. The Orchids - Natural History and Classification. Smithsonian Institution, Washington D.C., Chapters 1-3. Faletra, P., A. Dovholuk, K. Sokolski & T. King. 1997. Saving Cypripedium reginae. Orchids 66(2): 138-143. Hoshi, Yoshiikazu., K. Katsuhiko & H. Shuuichi. 1994. in Vitro Seed Germination of Four Taxa of Cypripedium Lindleyana 9(2): 93-97. Scagel, Robert F. et. al. 1965. An Evolutionary Survey of the Plant Kingdom. Wadsworth Publishing Company, Inc. Belmont, California. pp. 584. Sokolski, K. & P. Faletra. 1997. Growth Studies of the Showy Lady‟s-slipper (Cypripedium reginae) in Axenic Culture. Bulletin of American Association for the Advancement of Sciences , Annual Meeting ,1997. Withner, Carl L. 1974. The Orchids Scientific Studies. John Wiley and Sons Inc., New York.

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ALL TRESSED UP, BUT NO PLACE TO GO or Local Ladies’-tresses Orchids of Ohio Thomas A. Sampliner
Editor’s note: Botanical keys are often the bane of the amateur and professional alike. In all of our native orchids no genus has more similar species than the genus Spiranthes. In preparation for a field trip in northern Ohio, Tom had prepared a comparison of several books and field guides. I asked him to expand it to make these comparisons more pertinent for many of our members. It was not practical to reprint the various keys here, but if you have any or all of the books it would certainly be helpful to lay them out and follow along with Tom‟s comparisons. Please keep in mind that regional books cite habitats and flowering dates for their specific areas and often that data does not strictly apply elsewhere.

At various points, most of the following authors have had to fudge their keys by combining many traits just to make one dichotomous choice. This represents not only a weakness of a key for practical use but also violates the concept of what a key is supposed to be. An even greater failure of some of the keys occurs when the author not only had to combine many traits

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to make a separation between species, but had to require further choices between groupings of opposing traits before you can separate out another species. Careful forethought in using the traits is necessary to construct a key that will only require choices among one, or maybe two, opposing traits at a time which immediately separates out species. Admittedly, this is the ideal. You can judge for yourself how well each author has done. It must be stated that to have these experts manifesting conflicts in dividing small from large florets based upon lip measurement is inexcusable. The failure of such a trait is demonstrated by the authors‟ employment of measurements that not only conflict with each other, but also require exactness to the millimeter while in the field. If the experts cannot agree upon, or are not familiar with, existing writing so they cannot eliminate such measurements or explain to the lay person how measurements play a valid role in a key, then the trait is invalid. There were also some traits used that the authors felt necessary to caveat. This also is an inappropriate key trait. According to Carlyle A. Luer, in his monumental effort entitled The Native Orchids of the U. S. & Canada, excluding Florida (1975), the genus Spiranthes (sensu latu)3 consists of over 300 species
3

Current taxonomic thinking has considerably revised the genus so that there are 40-45 species and the remaining 300 or so are in a number of segregate genera. This makes the genus on the whole much more workable.

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ranging from temperate to tropical regions. Few species occur in the old world. Only one species each occurs in Japan, Australia and New Zealand. Africa, as of 1975, was not known to have any. In addition to Luer, the other written sources consulted for this article were: Frederick W. Case Jr.'s Orchids of the Western Great Lakes Region (1987); R. T. Whiting & P. M. Catling's Orchids of Ontario (1986); Paul Martin Brown's A Field and Study Guide to the Orchids of New England & New York (1993)4; and Homoya‟s Orchids of Indiana (1995). I had contemplated using Merritt Lyndon Fernald's key in the 8th edition of Gray's Manual of Botany (1950). My decision not to use it was based in large part on (1) the great differences in coverage, and (2) the traits employed were so different that a fair comparison with the others was impossible. The initial discussion will contrast the keys from each work with an emphasis on their treatment of local species. This article is intended to be a preparation and quick reference source for fall field trips in the Ohio area when it might be possible to see as many local species as a one day field trip might permit. Depending on how the growing season has progressed and the amount of driving you are willing to do in one
See Garay, L. 1980. A generic revision of the Spiranthinae. Botanical Museum Leaflets 28(4). Harvard University, Cambridge. ed. 4 Recently republished ( March 1997) in a totally new format with 192 color plates and new drawings as Wild Orchids of the Northeastern United States. Cornell University Press, Ithaca, NY. ed.

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day, you can see six or seven species dawn to dusk. You can see even more if additional days are available. It should also be useful for appropriate species in the entire northeast. Sadly, I must note that fall is too late in the year to see shining ladies’-tresses, Spiranthes lucida. In Ohio this species appears in June. Depending upon when in the fall you reach Ohio, the same may be true of northern slender ladies’-tresses, S. lacera var. lacera, southern slender ladies’-tresses, S. lacera var. gracilis and grass-leaved ladies’-tresses, S. vernalis. You should be able to find the other species in the northeast in accordance with local expected bloom periods. Whiting & Catling have selected lip size as the initial feature for their dichotomous key. The choice is between florets exhibiting lip size 4-7 mm in length versus 8-11 mm, with an additional trait for some species in the first category of being loosely spiraled while all under the second choice are to be densely flowered. Therefore, in our area, Case’s ladies’-tresses, Spiranthes casei var. casei, S. lucida, northern oval ladies’-tresses, S. ovalis var. erostellata, and both S. 1acera var. lacera and S. lacera var. gracilis would fall within the first division. It should be noted that as of their writing, little ladies’-tresses, S. tuberosa, had not yet been found in Ontario.

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Case uses lip size for his first choice as well. However, he opposes lips less than 6 mm long versus those 6-11 mm. No companion trait accompanies his first division. Case does include Spiranthes tuberosa in his key. Homoya starts his key by placing stems slender with small flowers, the lip of the lowest florets 5 mm or less, in opposition to stems stout with larger flowers, the lip of the lowest florets 7 to 10 mm. His key, therefore, also uses lip sizes as a means of differentiating species. Luer covers the widest geographical and climatological areas so his first dichotomy uses such traits as spiral appearance, ranking of the spirals, the lip growth (called a tuberosity), and climate. Obviously, this was done to separate the tropical, subtropical, southern, and western US species. We will, therefore, pass over these features and proceed to the first couplet relevant to the Great Lakes area. This is an examination of the spiral feature. He opposes secund (appearing one-sided) to conspicuously coiled, florets appearing in one rank versus tightly spiraled flowers appearing in more than one rank. However, he has a caveat under both choices that occasional specimens can appear under the other heading.

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Paul Martin Brown, in his key, has leaves narrow and grass-like at flowering time versus leaves orbicular or absent at flowering. For the novice orchidist, this is by far the easiest observation to use. It has easily observable characters that the other keys lack at this juncture. Based upon the practicality of using a key in the field in any region, but especially ours, I note a real problem with Luer's key and an apparent conflict between the size traits as used by Whiting & Catling versus Fred Case‟s. I do not mind trying to follow Luer in his spiral patterns, but if he finds a need to caveat the separating traits, how useful is it? Keys that conflict as to how to separate lip size are a real problem. I do not see how Case‟s 6 mm versus larger can be helpful if Whiting & Catling can distinguish lowest florets of an inflorescence of 4-7 mm versus 811. Homoya also uses lip size initially, but he provides a very practical application by combining size with the easily observable contrast of slender versus stout stems. No one else employs this trait. His lip measurements also avoid the dilemma posed by the conflicting lip measurements in the keys of Case versus that of Whiting & Catling. Several of the authors employ ranking as a trait key. Homoya does as well, but he goes further than the others by adding the concept of phyllotaxy to it. Here Homoya cites Sheviak (Biosytematic Study of the Spiranthes cernua Complex, 1982) explaining that this is the number of flowers necessary in the spiral to place the next

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possible floret in a direct line with the one from which you started. This is then expressed as a fraction, so that if it took 5 flowers to get back into perfect alignment, then the phyllotaxy is 1/5. Homoya excels in his ability to communicate various botanical concepts in clear yet precise language. So that we can more clearly discuss the keys, I will borrow from Homoya to set forth ranking and phyllotaxy of the spiral florets. Ranking is the spiral arrangement of the florets which can be discernible as single, if so loose that a staircase effect appears; or multiple to indiscernible, when florets are so crowded and tight that two, three, or more separate stairways of florets appear when you are looking up or down the spike. If you look from the top of the spike down, you can determine the phyllotaxy. Homoya contends this concept is more useful than ranking in separating confusing or atypical specimens. Certainly this is observable in the field and it avoids the caveats regarding spiral traits found in several keys. I must say, I find Brown's first trait the most highly visible compared to the others, with Homoya a close second. Shape of and presence or absence of leaves is readily visible in the field! Let's continue to see more of who wrote what and how well they serve our purpose when we are in the field. Under the small floret species, Whiting & Catling‟s key uses rostellum and viscidium lacking,

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cauline (stem) bracts with long, spreading, recurved blades versus the parts present and bracts being bladeless. Choice one keys out to Spiranthes ovalis var. erostellata. The other choice requires further trait selection for species we will not see on a single field trip in the month of September in the Cleveland area. This key is of limited value on the small-flowered species, unless we find either of the varieties of S. lacera or S. casei. Frankly, even if we did find these species, how practical for field use are such esoteric features? Case, under small flowered species, next uses lip color: floor of the lip with bright yellow or green in the center versus white or pale, creamy yellow throughout the lip with no strong central coloration. This is usable. Under lip color he has the two varieties of Spiranthes lacera as well as the June blooming S. lucida. Under the white color we only find S. tuberosa. This latter species is so distinctive by size alone that either way we finish up with the locally expected species. Brown's key ignored size and started with leaves. Next he uses bright yellow central portions to the lip, taking you immediately to Spiranthes lucida. Lip otherwise sends you next to flowers white versus otherwise. Other than white florets sends you to the same point in his key that the 2nd choice for his first trait sent us (leaves orbicular or absent at flowering time). Therefore, Brown's key will lump: S. lucida,

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hooded ladies’-tresses, S. romanzoffiana, giant ladies’-tresses, S. praecox, and nodding ladies’tresses, S. cernua, as all having leaves narrow and grass-like at flower time; S. lucida keys out at strong yellow in lip central; fiddle (pandurate) shape lip keys out to S. romanzoffiana; and multiple ranking keys out to S. cernua from the S. praecox as well as their habitat differences. Very usable in the field!— especially since S. cernua is quickly removed from yellow ladies’-tresses, S. ochroleuca and S. casei with which it can be confused. Luer's third key trait under the small flowered species contrasts leaves that are narrow or linear, lanceolate to oblanceolate versus leaves ovate, but often fugacious (withering early). This is a useful trait for field work. Just to list the species that fall under linear-like leaves shows how useful: Spiranthes vernalis, S. praecox, cienegas ladies’-tresses, S. delitescens (as S. graminea), lace-lipped ladies’tresses, S. laciniata, S. casei ( as S. intermedia), longlipped ladies’-tresses, S. longilabris, S. ochroleuca and southern ladies’-tresses, S. torta (syn. S. tortilis). Sorting through what we may find here in Ohio, this separation is very helpful. Assuming that S. vernalis is still possible, S. ochroleuca is the only other likely local species and lip color in S. vernalis easily will separate these two. Since S. ochroleuca is one of those that is in the original S. cernua complex,

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it is extremely valuable if we can depend upon fugacious leaves to quickly sort out S. ochroleuca. Homoya next contrasts bloom time combined with lip color. His first division is flowering May to June, with yellow lip, which immediately keys out to Spiranthes lucida. The contrast has flowering time July or later combined with lip green or white. At first, I didn't think the category was contrived; however, I notice he had three small floret species left; namely, Spiranthes lacera, S. tuberosa, and S. ovalis. To progress, he contrasts under the late bloomers, green versus white florets to separate out S. lacera. That leaves two late blooming small floret flowers which viewed in isolation could be difficult for the amateur. He employs his phyllotaxy concept so that S. tuberosa keys out with a 1/5 versus the 1/3 to 1/4 of S. ovalis. He does also use lip shape and leaves absent or only basal for S. tuberosa while S. ovalis has the conspicuous stem leaves during bloom time. A nifty solution for the final small floret species. Next, let us look at the larger flowered species. In order to get to Luer‟s treatment of the larger floret species, we first looked at the spiral traits; then under tight spiral, multi-ranked if we select narrow leaves, we find a choice for lip size. Luer divides the species from those under 9 mm to those above. Just what we needed! Remember Case versus Whiting & Catling? I can see it in the field: is it between 4-7 mm? more or

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less than 6 mm? or more or less than 9 mm? If the experts can't agree what can we do? If we can get beyond this problem, continuing with Luer, his under 9 mm next requires selection for lip color and then habitat, including only Spiranthes ovalis among species we are likely to find. Therefore, we have a clear shot by using his larger than 9 mm lip to find our local species. He next opposes lip constricted near the center and northern or northeastern distribution versus lip ovate and usually not constricted near the center. If the lip is oblong to ovate and only slightly constricted near the center you are left with S. cernua or S. ochroleuca. If the lip is pandurate (fiddle-shaped), dilated apex and white flowered you have S. romanzoffiana (or in Luer a cross called S. xsteigeri, which has proven to be S. ochroleuca). In our area we need be concerned only with the first two. Spiranthes cernua will have the lip base dilated, tuberosities small and flowers uniformly all white. Spiranthes ochroleuca will not be dilated at the base, tuberosities large and flowers will have a yellowish cast. Back to Whiting & Catling for their larger flowered species. Here, the first choice has basal portion of the rachis glabrous (smooth), hairs sparse and only 1 mm long: lateral sepals 3 mm or wider, lip pandurate, perianth veins prominent; which immediately identifies Spiranthes romanzoffiana. The opposing features would be: basal portion of rachis

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pubescent with capitate hairs 0.3 to 0.5 mm, lateral sepals 2-3 mm wide, lip not pandurate, perianth veins not prominent. Next, under the second choice, we select between: leaves absent at flowering, cauline bracts 4-7 mm, flowers white but central portion of lip is yellowish and thickened, flowers fragrant, lateral sepals diverging, calli (small, thickened calluses) 0.5 mm long which keys out Great Plains ladies’-tresses, Spiranthes magnicamporum. The alternative is: leaves present, cauline bracts 2-4, flowers white or cream color, no strong fragrance, lateral sepals appressed, calli 0.7-1.5 mm. This requires a further choice to separate S. ochroleuca from S. cernua. If flowers are cream color, dorsal and ventral curvature of the flower pronounced, measuring a separation of between 0.5 to 1.5 mm between dorsal and lateral sepals, lip thick, fleshy centrally, often with slight basal dilation you have Spiranthes ochroleuca. The opposite choice requires white flowers, moderate curvature resulting in dorsal from lateral sepal separation of 0.1 to 0.9 mm, lip neither thick nor centrally fleshy, and without distinct lip dilation you are led to Spiranthes cernua. Rather than looking for these features, which are somewhat difficult in the field, I find helpful a habitat observation whereby S. ochroleuca will be in the sandier more well drained site. This can sometimes be easily seen above the wetter area, including ditches, which are preferred by S. cernua. Frankly, considering

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all the keys‟ attempts to separate these two species, this is probably the most useful and user-friendly observation for field work. Case classifies large-flowered species as lip 611 mm. The first choice is lip pandurate, small basal callosites, sepals and petals connivent forming a hood, inflorescence heavy and thick, not strongly spiraled, secund in rank. This keys to Spiranthes romanzoffiana. The opposite is lip not pandurate, has prominent basal callosities, no hood, strongly spiraled spikes. Under this latter choice, the next selection in the key will direct you to Spiranthes magnicamporum with leaves absent at flowering, lip deflexed giving a gaping appearance, central lower lip yellow, above thick white, callosities short, conical, as wide as high, lateral sepals free and spreading, flowers strongly fragrant. Its opposite is: leaves present at flowering, lip color uniformly white or creamy, basal callosities longer than wide, lateral sepals appressed. Under this latter category we elect between: flowers pale cream, yellow or faint green tone, upper lip surface deeper yellowish cream versus flowers white, crystalline and in multiple ranks, lip white throughout, dorsal and lateral sepals and petals approximate, slightly recurved but gaping or a 2-lipped aspect, leaves present in good condition at flowering. These criteria immediately identify Spiranthes cernua.

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Under the first choice above, he separates the last two: Spiranthes casei from S. ochroleuca. Spiranthes casei is lip 6-7 mm dorsal sepal and lateral petals directed forward, scarcely upcurved, lateral sepals forward, flowers spaced so that groups appear as florets of 2-3 in loose spiral; lower leaves shriveled, upper-most persist at flowering. Finally, S. ochroleuca under Case‟s key has lip 7-10 mm, dorsal sepal and lateral petals connivent, recurved providing a 2-lipped appearance, lateral sepals ascending, prominent basal lip callosities, lowermost leaves present in good condition at flowering. We left Brown‟s key at couplet #6. The choice is flowers with green central portion versus not. The aforementioned leads to #7, wherein Spiranthes lacera var. lacera is identified with loose spike, few twists and those near the spike summit, basal leaves present at flowering. Spiranthes lacera var. gracilis calls for dense florets on the spike, regular twisting on the spike, basal leaves withered at flowering. Without green central lip pushes us to #8 wherein flowers purest white is Spiranthes tuberosa versus pale cream, yellowish white or bicolor. This moves us onward to choice #9. Here flowers yellowish/white keys to S. ochroleuca. Flowers white with yellow/beige lip leads to choice #10. This is the last couplet: flowers slender, never fully expanded, plants of northern areas identifies as S. casei. Flowers

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fully expanded, lip contrasting to petals and sepals identifies S. vernalis. Homoya starts his large-floreted species with a conglomeration of four widely divergent traits. It must be said that keys should oppose one, maybe two traits at each selection point, not up into the four or more range. The more combined choices the more unwieldy the key. His first selection requires: single rank spiral of 1/5 or smaller phyllotaxy; sepals inrolled toward the base; pubescence pointed; plants of dry grassland; flowering season from mid-June to July. The combination is said to isolate Spiranthes vernalis. In opposition: multi-ranked, phyllotaxy of 1/3 or 1/4; capitate pubescence; sepals flattened at the base; blooms September or October with a caveat that it is even earlier in northern wetlands. I hope you agree that this is not an effective key since this selection requires further choices to key out any further species. Therefore, you must next isolate: lip pandurate; ascending flower parts; petals and sepals joined to form a hood which together key out to Spiranthes romanzoffiana. At this point you still need to make further choices under the opposing traits before you get another species. Thus you must look for: lateral sepals widely spreading with tips arching above the petals; fragrant flowers, leaves fugacious and, he claims, calcareous habitat in either dry grassland or

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dunes for S. magnicamporum. This may be correct for Indiana but definitely not for Ohio. Ours like wet, disturbed areas as in the Cuyahoga Valley National Recreation Area amidst a glaciated plateau which was disturbed during freeway and other construction and is mostly connected with borrow pits. Novices could be confused if Homoya‟s key were followed religiously. Opposing traits in this key bring you to the final selection which pits the traits of Spiranthes ochroleuca against those of S. cernua. For the former, seek lateral sepals appressed and parallel to petals but ascending with lip curvature obvious; flowers cream to yellowish; basal tuberosities prominent. Spiranthes cernua presents whiter flowers, less prominent tubercles and a wide assortment of habitats. As previously mentioned, the observation by Whiting & Catling, and also Case, that given a wet ditch, S. cernua likes to be down toward the wet, while S. ochroleuca will be higher and drier, is the most useful field trait. There are obvious strengths and weaknesses in each key. Comparisons are difficult but some I've shared with you already. Since we are Ohioans, I should mention in passing that due to limited species recognized at their times of publication there is no point in trying to use either Clara G. Weishaupt's Vascular Plants of Ohio (1971) or E. Lucy Braun's The Monocotyledoneae (1967). The only species covered are: Spiranthes vernalis, S. gracilis, S.

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tuberosa, S. romanzoffiana, S. lucida, S. cernua and S. ovalis.

CONCLUSION: Happy hunting and may you now feel that you will no longer have any trouble with the tresses; come bloom season they will be awaiting you, for they are all tressed up with no place to go.
THOMAS A. SAMPLINER

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WILD ORCHIDS OF THE NORTHEASTERN UNITED STATES
A Field Guide by Paul Martin Brown, drawings by Stan Folsom Comstock Publishing Associates a division of Cornell University Press, Ithaca and London. 1997. 236 pp.; 48 pp. color plates. $17.95 paper. ISBN 0-8014-83417. The last 15 to 20 years have witnessed the opening of the golden age of the regional field guides. Regional works have much to recommend them: a limited geographic area typically limits the number of species included allowing for more in-depth treatment, and authorship by someone with extensive experience within that area assures a level of practicality of information difficult to achieve in a national guide. Wild Orchids of the Northeastern United States, a field guide written by Paul Martin Brown and illustrated by Stan Folsom, incorporates all the positive features of a regional guide to make a significant contribution to the botanical literature of the northeastern United States. Some of the special strengths of this work are the completeness of the species listings, the accuracy of the local flowering seasons, the author's appraisal of the current status of each orchid as reflected in the distribution maps, and frequency of occurrence comments, and the state by state checklists. Even

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experienced orchidologists will find valuable insights into little known species such as Epipactis atrorubens and Gymnadenia conopsea. I found the distribution maps to be very accurate, reflecting both current species distribution and important historical knowledge. By combining these maps with frequency comments and state checklists, the reader can easily form an impression of the status of any species in a given state. Numerous (192) color photographs contribute to the beauty and depth of this volume, as they depict both the typical and variant forms of the orchids. Stan Folsom's line drawings attractively and accurately depict each species in this book. If I were to make one suggestion for future editions, it would be that whenever possible these drawings be life-size, or if not then the scale of enlargement or reduction be included. Wild Orchids measures nine by six inches and is 236 pages long, a convenient size to carry afield or to "plant" on a personal library shelf. Everyone from wildflower enthusiasts to orchid experts will want to add this volume to their collection.
William K. Chapman, PO Box 184, New Hartford, NY 13413. Bill is the author of Wild Orchids of the Northeast reviewed in this issue, and numerous other natural history books.

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ORCHIDS OF THE NORTHEAST
by William K. Chapman Syracuse University Press, Syracuse, NY 1997. 195 pp. $32.95 ISBN 0-8156-2697--5 cloth ; $17.95 ISBN 08156-0342-8 paper Spring of 1997 appears to have not only brought the usual array of wild orchids (and other wild flowers, too), but also a wave o local native orchid field guides and publications. My current book (Wild Orchids of the Northeastern United States) and Bill‟s have been running parallel for two years now and, as we both predicted, have become available within a few weeks of each other. In planning review I thought it only fitting that we should review each others work. For those who do not know us we have spent many hours together in the field and it the two field guides were not planned to appear together - it just happened! Bill Chapman‟s work present a very different aspect of a field guide with extensive text relating to the historical information of the species treated and even more information regarding floral measurements and personal observations. Each species is accompanied by a color photograph inset and keys are provided to each multi-species genera as well as to the entire treatment. The key are innovative and work well in the field. Chapman‟s treatment of both the Cypripedium parviflorum complex and The Platanthera huronensis-hyperborea complex are

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extensive and most welcomed. Any more information that helps local orchid enthusiast sort out these often confusing groups is always a major plus. General ranges are given for each species and an extensive glossary and bibliography. In comparing it to my book the complementary aspects would include the section on entymology for each species and the detailed information on measurements. As both Bill and I have known all along the two field guide do complement each other and together present the most complete information, photographs distribution maps and drawings available for the orchids of northeastern United States. (PMB)

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THE ORCHIDS IN THE OTTAWA DISTRICT: FLORISTICS, PHYTOGEOGRAPHY, POPULATION STUDIES AND HISTORICAL REVIEW by Joyce M. Reddoch and Allan H. Reddoch Special Issue of The Canadian Field-Naturalist VOLUME 111, Number 1, 1997. CAN$10.00 & $2.50 shipping.
This 186-page work describes the 44 orchid species that have been found within 50 km of Canada's National Parliament Buildings in Ottawa. It contains information on identification, past abundance, population changes, development cycles and relative stability of colonies. It is presented as a baseline study from which to design further research and prepare effective planning measures to protect wild orchid populations. The Introduction describes the history of collecting and recording since 1856, principal orchid habitats, local distribution patterns, rare species, colour forms, capsules and seeds, blooming dates and other topics. Each species account provides detailed information on the above topics, as well as a brief description of the plant. A drawing and a spot distribution map accompany each account. Correlations of some species with the Canadian Shield or the St. Lawrence Lowlands, or with calcareous rock, sandstone or sand deposits are shown. Longlived colonies of many species are described, and population studies are included for Corallorhiza striata, Goodyera pubescens, G. tesselata, Platanthera hookeri, P. orbiculata and Spiranthes cernua.

With 30 years of field observations amassed the Reddoch‟s have assembled what is perhaps more information on a local area than any other local orchid

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flora. In most instance local floras of such a limited area have information that is basically of local interest. Not so in the publication! Without question this is one of the most meticulously executed local orchid floras to every be published in North America. In addition to all of the botanical information and data, it contains a vast mount of local natural history. The line-drawing illustrations by Susan Laurie-Bourque are exquisite and in great detail. Several species also have fruiting spikes illustrated. This is to be highly recommend for all North American native orchids enthusiasts. Because it is a regular issue of a journal the run may be limited so do not hesitate to order yours immediately. (PMB) To obtain copies of this journal issue, send CAN $10.00 plus $2.50 (postage and handling) for each copy to: The Canadian Field-Naturalist PO Box 35069, Westgate PO OTTAWA, CANADA K1Z 1A2

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HIDDEN ORCHIDS
Photography by Thomas J. Bulat, Text by Thomas J. Bulat and Marilyn Bulat Rudi Publishing, San Francisco and Iowa City 112pp. ISBN 0-945213-20-4 ISBN 0-945312-19-0 paper. 1995. $44.95 hardcover $32.95 paper The subtitle of this large-format volume is “a photographic discovery of the disappearing native orchids of the United States and Canada” To my knowledge this is the first large-format photograph album of native orchids to become available. Thirtyfive species, primarily from eastern North America, are presented, some with several different views. The color is lavish and the angles often very dramatic. Often an extreme close-up is accompanied by a line drawing of actually size. This is most helpful. The accompanying text frequently includes a poem by Mrs. Bulat and narrative or field notes by Dr. Bulat. It is certainly a welcome addition to the North American orchid literature and affords an opportunity to view some dramatic photography. A few of the photos are ambiguously identified. Plate 2 - purple fringed orchid is large purple fringed orchid, Platanthera grandiflora; Plate 3 small yellow lady‟s-slipper is Cypripedium parviflorum var. makasin; plate 4 is eastern fairyslipper, Calypso bulbosa var. americana; Plate 29 &

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30 is Great Plains Ladies-tresses, Spiranthes magnicamporum, not nodding ladies‟-tresses, S. cernua; plate 49 & 50 is western prairie fringed orchid, Platanthera praeclara. These amendments in no way detract from the beauty of the work and in most cases are clarifications and/or species which have been described in the past twenty years and do not appear in all of the literature. Appendix A - Identification, Location and Conservation, presents good basic information on its subjects but contains the one most often perpetuated errors in so many native orchid articles and books It states that epiphytic species are confined to southern Florida. This is not true as several species range northward to southeastern North Carolina and westward through Louisiana. Most notable and wideranging is the green-fly orchis, Epidendrum conopseum. Appendix B is an annotated list of the photographs with more details on their locales and notes the sizes of the plants. Although the book was not widely distributed upon publication we urge members to secure a copy while they are still available. (PMB)

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