June 2007 North American Native Orchid Journal

NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 13 (2) 2007

IN THIS ISSUE: A Synopsis of TWO NEW PLATANTHERA SPECIES FROM THE WESTERN UNITED STATES DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS: CYPRIPEDIUM CANDIDUM GOOD THINGS COME IN SMALL PACKAGES or TINY JEWELS IN THE WILD

and more………….

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

NORTH AMERICAN NATIVE ORCHID JOURNAL

Volume 13 (2) 2007

CONTENTS NOTES FROM THE EDITOR

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Looking Forward October 2007

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A Synopsis of TWO NEW PLATANTHERA SPECIES FROM THE

WESTERN UNITED STATES Paul Martin Brown

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DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS:

CYPRIPEDIUM CANDIDUM Margaret M. From

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GOOD THINGS COME IN SMALL PACKAGES or TINY JEWELS IN THE WILD

The Slow Empiricist 75

MY FAVORITE THINGS A gallery of orchid photos

79 THE EFFECTS OF VARIOUS MEDIA AND ADDITIVES ON THE GERMINATION AND DEVELOPMENT OF SEVERAL NORTH AMERICAN NATIVE ORCHIDS

David Niemann 85

Note of Interest:

Epipactis palustris - Another European Visitor New to the North American Orchid Flora

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TEXAS LADIES’ TRESSES (SPIRANTHES BREVILABRIS) REDISCOVERED IN TEXAS

Eric Keith 113

Book Reviews 115

Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey Orchids in Hawaii

Vanishing Beauty: Native Costa Rican Orchids 1. Orchid Arabia

Orchids of Europe, North Africa and the Middle East Orchidelarium

In Memoriam – T. Sanders McMillan IV (1974-2007) Jim Fowler

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ON THE REPORTS OF SPIRANTHES VERNALIS Engelmann & Gray FROM NEW MEXICO

William F. Jennings & Charles J. Sheviak 124

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

subject to peer review and popular articles will be examined for both accuracy and scientific content.

Volume 13(2) pages 59-126; issued July 5, 2007. Copyright 2007 by the North American Native Orchid Journal

Cover: Platanthera macrophylla by Stan Folsom

NOTES FROM THE EDITOR

As mentioned in the last issue what had been originally planned for a single March issue has become two issues. In addition to the articles that were extracted from the original March issue I have decided to continue the photo gallery in My Favorite Things as well as several more book reviews. In this issue David Niemann‘s lengthy article on germination and media of several natives will be of interest to the specialist and the remaining articles should have an appeal to a broader audience. Stan Folsom and I will be making a major trip to the Southwest this summer to photograph a few select species for our latest book in preparation: Wild Orchids of the Southwestern United States, but should be able to access email every few days. I am soliciting additional articles for both the March and October 2008 issues at this time. Please submit both ideas and finished work. I can work with you on your ideas if you are not quite sure how to develop them. Paul Martin Brown Editor [email protected]

LOOKING AHEAD……

OCTOBER 2007

THE NATIVE ORCHIDS OF COLORADO

by

SCOTT F. SMITH

Book Reviews

& Notices

Brown et al.: Synopsis of TWO NEW PLATANTHERA SPECIES

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A Synopsis of TWO NEW PLATANTHERA SPECIES FROM THE

WESTERN UNITED STATES

In a continuing saga of the green-flowered species of Platanthera in the western United States two new species have recently been published. Platanthera tescamnis Sheviak & Jennings, the Intermountain rein orchid, in Rhodora 108: 19-31, 2006 and Platanthera yosemitensis Colwell, Sheviak, & Moore, the Yosemite bog orchid, in Madrono 54(1): 86-93, 2007. Both of these are segregates from P. sparsiflora/stricta/hyperborea complex, a group that has previously yielded P. purpurascens (Rydberg) Sheviak & Jennings (North American Native Orchid Journal 3(4): 444-49. 1997) and P. zothecina (Higgins & Welsh) Kartesz & Gandhi (Great Basin Naturalist 46: 259. 1986). Due to publication limitations color photographs and/or distribution maps of both new species were not available in the original publications. The species have been synopsized here in field-guide style pages and much of the material has been synthesized from the original descriptions. The reader is referred to those original publications of all four species for full details regarding the technical aspects of the plants and specimen citations. I am very grateful to Alison Colwell, Peggy Moore, Chuck Sheviak, Bill Jennings, and Scott Smith for their help in gathering information to put these pages together. PMB Note: Madrono 54(1) Jan.-Mar. 2007 was published on July 3, 2007.


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Platanthera tescamnis Sheviak & Jennings

Intermountain rein orchid

A new Platanthera (Orchidaceae) from the Intermountain West. Rhodora 108: 19-31. 2006 Range: primarily the Great Basin and Colorado Plateau in western Colorado, Utah, Nevada, and rare in bordering areas in eastern California and northeastern Arizona Plants: 29-126 cm tall; leaves usually 4-6 near the base of the stem, lanceolate to 29 cm long, ascending to spreading, inflorescence 1/3 – ½ the heighth of the plant Flowers: green to greenish yellow; ca. 1 cm long, the lip greenish yellow, linear-oblong to oblong, varying to lanceolate at the western margin of the range, to 7 mm long, the spur generally pendant, slightly clavate or cylindrical to 7 mm long; proportion of spur to lip is 4/5 – 1.2/5; size of the column—small and narrow—and position of the pollinia—close to the column—is distinctly different from other western species of green-flowered Platanthera. Habitat: along waterways in the drier areas; the name meaning desert (tesca) and swift-flowing river (amnis). As with most of the western species of Platanthera habitat can be variable but this species is distinctive for its preference for essentially mesic to dry areas near seasonal wetlands at elevations varying from 1825-2950 m Flowering time: late June to early August

Platanthera tescamnis, the Intermountain rein orchid, has been long known in the region and usually identified as P. sparsiflora. Although P. sparsiflora occurs elsewhere in California, Arizona, Utah, and Nevada, with the description of P. tescamnis, P. sparsiflora it is not known from Colorado. Plants of the ‗hanging gardens‘ from the four corners area are P. zothecina. Platanthera tescamnis preference for drier areas and at lower elevations than other related species often aids in identification. Only in the eastern limit of the range in Colorado does it occur with P. aquilonis. documented sites reports


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Platanthera tescamnis, Colorado

photos by Scott Smith


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Platanthera yosemitensis Colwell, Sheviak, & Moore

Yosemite bog orchid

A new Platanthera (Orchidaceae) from Yosemite National Park, California. Madrono 54 (1): 86-93. 2007

Range: Mariposa County, California; currently known from few sites a very restricted area in Yosemite National Park Plants: 20-80 cm tall; leaves 5-7 on the lower 2/5 of the stem; lanceolate, strongly ascending and reduced to bracts Flowers: 30-50, loosely flowered with floral bracts about the same length as the ovary; sepals green to yellowish-green; petals and lip yellow; lip rhombic-lanceolate somewhat dilated at the base with the tip usually caught in the dorsal sepal but if free the lip horizontal, not descending; flowers and fruit diminishing in size upwards and the withered flowers remaining on the ripening capsules Habitat: wet, montane meadows and seeps usually in tall(er) grasses and sedges; often found along turfs and streamlets at an elevation of 2100–2285 meters Flowering time: July/August

Unquestionably one of the most restricted species of Platanthera in this section to have been found. Plants were originally collected as Habenaria hyperborea by G.H. Grinnell in 1923 and then rediscovered by Coleman and Glicenstein in 1993 and a fragmentary specimen was identified as P. purpurascens. This placed the latter species well west of all known sites for P. purpurascens. Subsequent investigation has shown that the Yosemite plants vary in several critical aspects from other species of Platanthera that are found in the region. Habitat (slender, open inflorescence) color (yellow corolla) fragrance (musky) and pollination mechanics all are distinctive in this species. In addition, the restricted range and specific habitat within that range raises questions as to whether additional sites will be found.


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Platanthera yosemitensis, Mariposa Co., California photos by Peggy Moore

From: DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS: CYPRIPEDIUM CANDIDUM

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DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS:

CYPRIPEDIUM CANDIDUM

Margaret M. From The small white lady‘s slipper, Cypripedium candidum, is a rare sight in Nebraska. The shy, diminutive orchid is found in only seven locations scattered widely across the state, and although some populations may have a hundred or more blooming individuals, species protection is critical for its long-term survival. The specific name candidum comes from the Latin word candere, which means ―to shine or to glow‖. In the sixteenth century lady slipper orchids were called ―Slipper of Our Lady‖, which was meant to pay homage to the mother of Christ (Simon, 1975). The conversion of tall-grass prairies to productive cropland over the last one hundred years left little more than a few native prairie remnants intact, so that now the encounter of even one orchid in bloom is a gem not soon forgotten. Populations in the state represent some of the most western locations within the range of Cypripedium candidum, making the genotype(s) important conservation subjects. Orchids face a precarious existence in the harsh climate of the Great Plains, where frequent droughts, bitterly cold, dry, winter winds and blazing summer temperatures test a species‘ mettle. Drought hit the plains particularly hard in recent years, and much of Nebraska has now experienced as much as 8 years of precipitation shortfall. That the orchids survive at all, under such stress, is simply amazing. Herbivores like to browse the orchids, often removing the tops before they can produce mature seeds to ensure a succeeding generation. Human safety takes precedence in road maintenance, sometimes permanently altering the orchids‘ habitat, as happened for C. candidum in Nebraska in 2005. When orchids are


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displaced by human activities, rescue operations are often the only option left; even if not the ideal. The human aspects are a fact of modern life, and will continue to put pressure on natural habitats. When an orchid population is faced with destruction what can orchid enthusiasts do? Some options are; to notify the appropriate governmental agencies, discuss with landowners how to protect the habitat, inquire about a conservation easement for the property, individuals or organizations can inquire about purchase of the site, one can move the orchids out of harm‘s way if permission is granted, or propagate and preserve some germplasm representative of the population so that the genetic diversity assumed present in the population does not disappear forever. Ideally, genetic studies should also be completed before any long-ranging decisions are made, but the reality is that destruction of habitat often happens with little or no warning, leaving no time to conduct all the research in advance. Two of the seven known Cypripedium candidum populations in Nebraska now face an uncertain future due to land sales and development, and a third faces environment-altering practices that may destroy the population. One of those populations is on the edge of a medium sized city where the orchid‘s habitat is scheduled for commercial development. All the sites have relatively good soil, which can easily be converted to crop production. The species as found in Nebraska, displays a range of variations, with some individuals possessing yellow sepals and petals with maroon-brown venations (Fig. 1), some having green sepals with maroon dots and venations, and some whose petals and sepals are entirely maroon-brown. Others have reported the maroon colorations as possible evidence of the relationship of Cypripedium candidum to Cypripedium parviflorum (Smith, 1993). An overlap of ranges for C. candidum and C. parviflorum is

believed to result in a natural hybrid known as Cypripedium xandrewsii Fuller. In Nebraska the nearest population of C. parviflorum is found within a woodland setting more than one hundred twenty five miles away from any of the sun loving C. candidum populations, although at some point in history the two ranges for the species may well have overlapped. Nebraska‘s C. parviflorum population was just discovered in


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the early 1990s, approximately 75 years after it was assumed extirpated from Nebraska, and botanists‘ early accounts of the state do not identify any overlap of the two species. The labellum is very similar in each of the variations. It is creamy white with light markings of magenta pink in lines along the sides and bright splotches of magenta on the inner rim, as well as on the lower portion of the column, near the edges. The leaves are reported as clasping the stem, according to Doherty (1997), but as the growing season progresses in the Great Plains, the leaves become fully expanded creating a lovely, open form (Figures 3, 5 and 6). Figure 1. C. candidum with mostly yellow-green petals and sepals.

Figure 2. Maroon petals and sepals on C. candidum


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Figure 3. C. candidum plants in protected habitat at the Omaha‘s Henry Doorly Zoo

In bud, the labellum is most prominent (Figure 4), and does not yet show the magenta markings that will be visible on the opened flowers later in the season.

Figure 4. A rescued plant in bud on May 6th, 2006.

As the blooming season comes to a close the flowers senesce and begin to take on a translucent appearance (Figures 5 and 6), but the leaves persist and appear to continue photosynthesizing for 3-4 months longer, producing energy that is translocated to the underground root


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structures where the flower and leaf meristems develop for the next growth cycle.

Figure 5. Cypripedium candidum as the flower senesces.

Rescued plants: Road work resulted in the displacement and damage to a number of the plants by the roadside at one of the native habitats. With permission of the state, some of the rescued plants were placed in a protected outdoor location and several others were potted up and placed under greenhouse conditions in order to study the relative merit of rescue operations if, over time, more plants face eminent threat of destruction. Rescued plants often die quickly after removal from their


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habitat (K. Kennedy, personal communication). The orchids were scraped from the roadside by road grader activity that damaged roots on most of them, and they were found covered with masses of debris at the bottom of the ditch (Figure 7). All rescued plants survived and 85% bloomed the first year, both the individuals planted outdoors as well as those that were potted up and grown under greenhouse conditions. It was decided to retain some of the native soil around the roots for those that were not bare-rooted when found and those specimens produced somewhat more vigorous growth over the growing season, as compared to those in the greenhouse.

Figure 6. A rescued plant as the flower begins to fade.

There are risks whenever orchids are removed from their natural environment, not least of which is that it is difficult to compensate for local conditions and adaptations, especially if plants are moved any great


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distance. Those local adaptations are another important reason that germplasm in the form of plants, seeds or pollen, should be preserved by long-term means if possible, so that if the need arises and good habitat is available, preferably in the original area, plants representing the local wild type could then be used for reintroduction.

Figure 7. C. candidum plant emerging from the roadside debris.

Seed-banking the local wild-type: Seeds are possibly the most efficient forms of germplasm preservation because they carry the possibility of genetic diversity and they are so tiny that great numbers of seeds can be stored within a very small space. Cryopreservation protocols are being worked out so that the diversity of Nebraska Cypripedium candidum populations will be represented and available for future conservation action plans. Until the time when efforts to keep wild populations safe and healthy are in practice, and the species‘ propagation and reintroduction methods are fully outlined, cryopreserved seeds provide materials for future research. Orchid seed cryopreservation protocols have to be researched on a species by species basis because each species has its own unique physical properties and culture requirements. The ability to preserve seeds for


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years, or even decades, is a powerful tool for conservation, provided that along with implementation of those protocols we also learn to better protect remnant habitats, understand the pollinators, and continue to study the microbial associations that are accepted as critical for the species. Seeds for C. candidum have already been tested in cryopreservation and trials are being conducted on their germinability in post-thaw cultures in the laboratory. The project is preserving seeds collected from the population facing the most immediate threat of destruction in the state. Some of the seed testae are damaged in cryopreservation tests (Figures 8 and 9), but other embryos are able to withstand the extreme temperatures of liquid nitrogen (-1960C), and still germinate. Results from the project will determine whether this is a feasible means to preserve C. candidum seeds for future conservation actions in the state. Figure 8. (left) Cryopreserved Platanthera sp. seeds, SEM at 194x.

Figure 9. (right) Scanning electron micrographs (SEM) at 372x, of cryopreserved tropical orchid seeds.

If practical it would be ideal if each state, province or region could preserve local genotypes for the long-range future, whether in cryopreservation or in parent populations grown under protected conditions. Keeping the populations numbers large is important, since plant species that maintain large populations stand a better chance of attracting pollinators and surviving over the long term. In order to


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maintain healthy populations there need to be increased incentives for habitat protection by private landowners, and research should be encouraged about pollination biology, edaphic changes, pollinator populations, mycorrhizal associations, soil conditions, and plant communities across the orchid‘s entire range in order to save the species.. Above all, those who have knowledge about Cypripedium candidum are encouraged to educate others in the region about the species itself, and about what can be done to save these beautiful members of our natural legacy. Acknowledgements: My thanks are extended to Jim Pyrzynski and Tim Janssen of the Greater Omaha Orchid Society for their assistance and perseverance to save the Nebraska Cypripedium candidum wild-type. Gratitude is also extended to Mike Fritz, biologist for the Nebraska Natural Heritage Species program at the Nebraska Game & Parks Commission, and to Dr. Kathryn Kennedy at the Center for Plant Conservation in St. Louis, for their help and valuable information. Dr. Simmons, director of Omaha‘s Henry Doorly Zoo, continues to be an inspiration to all of us who know him, and the motivating force behind conservation efforts for threatened species.

Literature cited: Doherty, J.W. 1997. The genus Cypripedium; a botanical and horticultural overview.

North American Native Orchid Journal 3(1): 5-120. Simon, H. 1975. The Private Lives of Orchids. J.B. Lippincott. Philadelphia, Penn. Smith, W.R. 1993. Orchids of Minnesota. University of Minnesota Press, Minneapolis,

Minn. Received on 4/23/07: As a postscript to the article about Cypripedium candidum rescued and planted at the zoo; I am happy to tell you that every one of them has reappeared above ground this spring. And most of them are displaying more ramets than last year. We are, of course, hoping that they will persist over the coming years. Margaret M. From, Plant Conservation Scientist, Omaha‘s Henry Doorly Zoo, 3701 S. 10th St., Omaha, NE 68107 [email protected]

Glossary:

cryopreservation: preservation of seeds or tissues at extremely low temperatures

which in effect is supposed to stop metabolism, so that cells don’t age

edaphic: plant communities influenced by the soils, not just the climate (i.e.

effects from sandy soils, acidic soils, or nutrient-poor soils)

germplasm: an organism’s DNA (i.e. seeds are stored DNA in a frozen seedbank)

meristems: clusters of cells at zones that will eventually produce shoots or roots

testae: seedcoat

Empiricist: GOOD THINGS COME IN SMALL PACKAGES

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above: Prosthechea pygmaea Wally Wilder left: Spiranthes eatonii right Spiranthes brevilabris P.M. Brown


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GOOD THINGS COME IN SMALL PACKAGES or TINY JEWELS IN THE WILD

The Slow Empiricist

This idea came to me when I heard a fairly new orchid enthusiast talk about seeing Prosthechea pygmaea. It has only one site that is known for it in southern Florida. The orchid grows in a large colony on one tree in the Fakahatchee Swamp. The flower is so very tiny that is hard to see even when the flower is fully developed. Out of flower most people would not know what it was. It has been described as being about one quarter of an inch overall. It is a nice medium shade of yellow but not so brilliant that it catches your eye when you might be passing by. As the flowers mature and the petals and sepals disappear as it sets seed and it would be nigh impossible to find the leafy plants in the tangle of the south Florida wilderness regions except for the fact that it has colonized this one tree with what appears to be a large single plant.

This experience coupled with the fact that I had been helping Paul Martin Brown with a project to document and count the extant populations of Mesadenus lucayanus, which has slender spikes of tiny bronze flowers, got me to thinking about the other elusive little gems that are out there. Now some of these tiny wonders might be larger to eye than the aforementioned orchids but their rarity makes them fit into this topic nicely because if there are only a few sites for this orchid to be found it should be accorded the same wonder status as the truly small sized orchids that tantalize the orchid enthusiast to find and enjoy them. There are many tiny wonders in the orchid world that grow all over the United States but Florida seems to have a monopoly on some of them. To name a few that fit this category you have some of the Spiranthes, especially S. floridana and S. brevilabris which seem to prefer central Florida for their habitat. Spiranthes eatonii grows in the Southeastern United States and there are several sites in Florida. All these Spiranthes share enough characteristics to put them in the small wonder category. Spiranthes eatonii has tiny little flowers that spiral up a stem that can be no larger than the lead in a mechanical pencil. Spiranthes brevilabris and S. floridana are so rare they are easy to miss so even though they don't officially fulfill the tiny wonder categories they belong in this section, at least to my way of thinking.


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Then you have a difficult orchid to even see when you are standing right next to it. Harrisella is a leafless epiphyte that often likes to grow on old fruit trees. It is small enough so you can hold the entire plant in the palm of your hand. The flowers are miniscule but the fruits are at least large enough so you can see them when you are standing a few feet away. They look like tiny little bells spraying off the gnarled branches of old orange trees that are one of their preferred hosts. Next, we have a family of tiny orchids that are very rare. The smallest in this group called Triphora are the ones labeled rickettii. They poke up from the forest floor only about a few inches or so and their tiny blooms are very easy to overlook. The next in size is craigheadii followed by trianthophora, which is much larger but has an erratic blooming cycle. If you live in more northerly climes you have all the twayblades to keep you searching. These little orchids have flowers that often look like little dancing men but this entire plant family is usually only a few inches high. Many of these orchids have a coloring that blends in with their habitat. When you find a colony, though, you will be richly rewarded by their whimsical appearance. Now there are more tiny wonders to find out there so don't be surprised that I didn't include your favorite or feel miffed because it was left out. Just be aware that there are a lot more that are just as hard to locate and just as wonderful when you do find them. This brings up the first order of business when you set out to locate a tiny wonder. The best of all possible worlds is to find someone who has already seen the plants and can lead you right to them. Failing that you would need accurate directions to be able to have a chance at locating many of the plants. Lastly, you need a persistent nature and a keen eye for observing these miniature gems that you would most likely walk right by if you weren't attuned to looking for them. Once you gain some familiarity with the plants' habits and their preferred habitat you might be lucky to encounter a new site for them in your ordinary scouting for other orchids. All it takes is for you to be sensitive to the possibilities that the plant may occur somewhere else in similar terrain. Finding these orchids is probably your first concern but just finding them, although a great and satisfying experience in itself, should not be your only goal. You need to learn to look at these orchids and begin to appreciate the delicate qualities they possess. This may mean getting down to their level and examining them with a hand lens. Seeing them magnified shows their beauty much more clearly. You can see the delicate parts that create the whole and you might even get to see such tiny parts as their glistening glands or shining hairs. While you are getting


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down and dirty don't forget to examine the lovely leaves if they are present at blooming times. Sometimes they are basal rosettes and sometimes they climb up the stems. Sometimes there is just a single leaf while other orchids have multiple leaves. Different orchids may have different characteristics even in the same genus. This means you have to do some homework if you are going to find these plants and recognize them for what they are. Reading about them in good orchid field guides or in journals or other scientific publications can prepare you enjoy your excursions in your quest of a particular orchid. Then when you encounter your quarry you will be rewarded with lots of different experiences. First you should feel exhilaration at having discovered the orchid's hiding place. Then you should begin to experience the wonder of this tiny creation for its color, delicacy, shape, scent if it has one, and so forth. These sensory experiences are what keep orchid enthusiasts out there looking and experiencing nature's bounty. By using all the aids you need to enjoy the plant from merely associating with it to really getting up close and personal with it, you should come away with a deeper understanding of those particular orchids. I wish you all happiness in your hunting and exquisite experiencing when you discover them. Your Slow Empiricist Harrisella porrecta Wally Wilder

MY FAVORITE THINGS

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MY FAVORITE THINGS A gallery of orchid photos

Calypso bulbosa var. americana Greg Alikas

June 6, 2004 Golden, Colorado Digital Konica-Minolta A2

MY FAVORITE THINGS

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Platanthera leucophaea Al Menk

June 2006, Michigan, Digital Nikon D100 with AF28-105 lens

MY FAVORITE THINGS

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Cypripedium reginae Tom Nelson

July 8, 2006 Bruce Peninsula, Ont. Pentax K1000 film

MY FAVORITE THINGS

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Triphora trianthophora var. trianthophora Wally Wilder

Marion Co, FL Digital Kodak

MY FAVORITE THINGS

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Sacoila lanceolata var. lanceolata Linda Cooper

May 2006 FL Digital Minolta Maxxum 7D SLR

MY FAVORITE THINGS

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Cypripedium parviflorum var. pubescens lacking anthocyans

Olin Karch April 28, 2004, Upper Buffalo Wilderness Area, Newton Co., AR Olympus

C2500L digital.

Neimann: I. EFFECTS OF VARIOUS MEDIA ON GRMINATION

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I. THE EFFECTS OF VARIOUS MEDIA ON THE GERMINATION AND DEVELOPMENT OF SEVERAL

NORTH AMERICAN NATIVE ORCHIDS

David Niemann

Introduction Land development and agricultural practices create many side effects, not the least of which is the destruction of the habitats of many of our rare native plants. The uncommon species are in especially serious danger because they often grow only in very special habitats (Niemann, 1986). Once these few habitats are destroyed, an ecotype or perhaps even the species itself may be lost forever. Native North American terrestrial orchids are particularly susceptible to this type of extinction because they frequently have very narrow ecological tolerances. There are 78 species of these terrestrial orchids listed in the 8th edition of Gray’s Manual of Botany for the northeastern United States. Many are already extinct or extremely rare in parts of their range. With regard to Cypripedium reginae, Fernald states that ―Plant liable to extinction through raids by nurserymen and would-be cultivators‖ (Fernald, 1950). Something must be done if these rare and beautiful plants are to be rescued from extinction. An intelligent and far-sighted approach to the problem would involve setting aside the remaining habitats of these and many other rare wildflowers so that they may continue to grow and reproduce under natural conditions. At the present time there are too few organizations or individuals willing or able to undertake this task. Another alternative would involve a thorough study of the ecological requirements of the mature and seedlings stages of the species most in danger. With adequate knowledge, suitable habitats could be protected in areas such as state parks. This is an extremely difficult and time-consuming task and it has not proved successful in the few cases in which it has been tried. A third possibility lies in the development of a completely artificial method of propagation (Niemann, D. 2001). If plants could be propagated in some way and allowed to reach nearly mature size, they could then be grown in outdoor habitats on protected land. The test of asymbiotic methods of seed propagation of these native North American terrestrial orchids is the purpose of this study.


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Materials and Methods

In speaking of the large number of orchid culture media, Withner (1959) stated, ―As yet there are no comparative data on all of these media on the same brood of seedlings…‖. It is unlikely that all of the media recommended in the literature are suitable for all orchid species. Since the development of a precisely suited nutrient medium requires a great deal of time, a test of many of the recommended media could give a more rapid indication of the nutritional requirements of some of these terrestrial orchids. A test of twenty of the more common media was set up (See appendix for media and formulation techniques. (Table 1 shows ion concentrations) I collected all of the seed used in this experiment. The seeds of Cypripedium pubescens and C. reginae were collected in my garden in Elmwood Park, Illinois on September 1, 1968. The capsules were still green at the time of harvest. Seeds of Calopogon tuberosus were collected in Lake County, Illinois on September 20, 1968. Plants of Spiranthes cernua were collected at Clark Junction, Indiana on September 20, 1968 and the plants were grown under artificial light until October 13, 1968 when the seeds were ripe and harvested. Ripened capsules of Pogonia ophioglossoides were collected on November 29, 1968 in McHenry County, Illinois. Soon after being collected, the seeds were removed from the capsules, air dried, and placed in closed jars containing calcium chloride. The jars were kept in a refrigerator at 4 degrees C. until the seeds were planted. The culture room consisted of a walk-in refrigerator with evaporating coils mounted in the ceiling. This presented a problem as cold air flowed down over the culture tables when the refrigeration unit was running. This problem was solved with a small fan in the growth chamber which maintained enough air circulation to give a relatively uniform 24 degrees C. +/- 2 degrees C. The culture tables had plywood surfaces painted with semi-gloss white enamel. The light source consisted of one General Electric 40 watt daylight fluorescent per light fixture. Typing paper was placed around the lamp to reduce the intensity. The fixtures were an industrial type with an enamel reflector. The fixtures were suspended so that the lamp was 12 inches from the surface of the table. This arrangement gave relatively uniform light intensity on each of the six tables. The light intensity varied from 180 to 320 foot candles as measured by a Weston model 603 light meter at the start of the experiment. Over most of the surfaces the intensity was 240 foot candles. A preliminary experiment showed that an intensity of 400 foot candles was not harmful while an intensity of 600 foot candles was too bright and caused chlorosis of the seedlings. The intensity of about 250 foot candles was chosen because of the


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preliminary experiment and previous reports of the necessity of low light intensities for seedlings (Northen, 1962). The seeds were sterilized in a 2% Clorox solution (5.25 % sodium hypochorite) according to Redlinger (1961). The seed was shaken vigorously for 10 minutes in a 1:50 solution of clorox:water. The seed was then rinsed twice in sterile distilled water. The seeds were sown in two ways. The seed of Spiranthes was easy to suspend in sterile distilled water. The seed of this species was planted with a dropper pipette. Two drops of seed suspension were placed in each tube. Due to the number of seeds per unit volume of water this amounted to about 150 seeds per culture tube. The seeds of the other species could not be suspended. They were planted by removing about 100 seeds from the sterilization vial with a small spatula. The seeds were then injected into a culture tube with a stream of sterile distilled water. With some practice, fairly uniform quantities of seed could be sown in each tube. Three readings were taken on each tube at 60 day intervals after planting. The readings were taken using the growth index developed by Curtis (described by Spoerl, 1948). In previous orchid research results were difficult to interpret because a uniform method of taking data was not used. Knudson and other early workers used measurements such as fresh and dry weight of seedlings, seedling color, protocorm diameter, total leaf and total root length. These characteristics may be all right for some purposes, but only one reading can be taken on a group of seedlings. The growth index is probably the best method of taking data because several readings can be taken on the same group of seedlings without destroying them. A major advantage is distinguishing degrees of development. Protocorm diameter is misleading because a large protocorm is as undeveloped as a small one. The growth index takes into account the developmental stage of a seedling. There are six stages in my modification of the index. These may be seen in Figure 1. The index is calculated by counting the number of seedlings at each stage in a culture tube. From this the percent of seedlings at each stage is calculated. Then the percent of seedlings at each stage is multiplied by the stage number. The sum of these products is the growth index for that tube. The data obtained in this manner are suitable for statistical analysis (Spoerl, 1948).


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Table 1.

Chemical Composition of the Media Medium Nutrient Ions mM/l

NO3 NH4 K PO4 Fe SO4 Ca Mg Cl Mn Sugar Conc. gm/liter

1 .78 0.0 5.0 1.9 .018 2.5 1.9 3.9 0.0 .022 maltose 20.0

2 4.36 0.0 1.78 .12 0.0 1.48 1.22 1.46 .87 .02 sucrose 20.0

3 12.2 3.79 6.32 1.44 .089 3.0 6.1 1.02 3.34 0.0 glucose 10.0

fructose 10.0

4 12.2 3.79 4.71 3.27 .893 3.72 6.1 1.02 0.0 0.0 sucrose 20.0

5 12.2 0.0 1.84 1.84 .006 1.02 8.26 1.02 2.17 0.0 sucrose 20.0

6 5.2 7.5 7.04 2.48 .137 4.84 1.94 1.02 0.0 .034 sucrose 20.0

7 9.44 5.53 6.15 2.2 .089 .54 0.0 .43 0.0 0.0 glucose 10.0

fructose 10.0

8 12.2 10.6 4.74 2.94 .67 6.52 6.1 1.22 0.0 0.0 glucose 10.0

fructose 10.0

9 11.0 0.0 9.16 .18 .007 .15 1.44 .14 .87 0.0 sucrose 20.0

10 12.2 7.57 1.84 1.84 .112 4.95 6.1 1.02 0.0 .034 sucrose 20.0

11 12.2 7.57 1.84 1.84 .112 4.95 6.1 1.02 0.0 .034 sucrose 20.0

12 4.23 2.75 .88 1.01 .146 1.06 1.48 1.06 0.0 0.0 sucrose 20.0

13 12.2 7.57 1.84 2.10 .267 4.80 4.0 1.02 0.0 0.0 sucrose 20.0

14 7.02 2.75 .88 .88 .016 1.06 2.13 1.06 0.0 0.0 glucose 10.0

15 15.32 3.12 2.62 2.04 0.0 1.02 6.1 1.46 0.0 0.0 glucose 2.0

sucrose 18.0

16 16.7 4.43 6.06 1.27 0.0 0.0 0.0 0.0 .43 0.0 sucrose 35.0

17 sucrose 5.0

18 no sugar

19 sucrose 5.0

20 no sugar

All media are in Appendix III of Withner (1959), except Ito (1955), Kano (1968) and Vacin and Went (1949)

Statistical Design

A 5 x 5 Latin Square was used for the location of the five species involved on the five tables used in this experiment. Within the species the tubes of the twenty media were arranged in a completely random manner using the table of random permutations of twenty in Fisher and Yates (1953). This design distributes the effect of light variation across the table and temperature variation with height because all five species are located once in each position


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across tables and once on each individual table. The ―F‖ test described by Snedicor and Cochran (1946) was used to determine significance. Dormancy Test A preliminary test showed that seed of at lease one of the species being tested germinated better after a cool period. Seeds of Calopogon tuberosus which had hardly enlarged in 6 months of culture at warm temperatures resumed development after a cool period. The preliminary test showed that seeds of Cypripedium reginae, C. pubescens, and Spiranthes did not react in this way. The reaction of Pogonia to this treatment was not tested in the preliminary experiment, but its behavior was similar to Calopogon in some other respects so it was included in the cool treatment. To further test the effects of a cold treatment, the tubes of Calopogon tuberosus and Pogonia ophioglossoides were placed in refrigeration at about 4 degrees C. with no light for 40 days following the 180 degree reading. Though Stoutamire (1964) makes no mention of this enhancing the germination, he suggests that 4 to 6 weeks of refrigeration are required to break the dormancy of the corms of Calopogon after they have been produced by seedlings. The preliminary experiment showed a cool period of 40 days to be very beneficial, and this is why that length of time was chosen. After removal from refrigeration, the tubes were replaced in their original positions on the tables and readings were taken after 60 additional days (the 240 day reading) using the growth index. Figure 1.

Stage 1 - dead seed or seedling Stage 2 - ungerminated seed Stage 3 - embryo breaking through seed coat Stage 4 - leaf point emerging Stage 5 - seedling with leaves Stage 6 - seedling with leaves and one or more roots


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RESULTS The results of the test of the nutrient media were surprising in several ways. The species effect was especially unexpected. Seeds of Cypripedium reginae and C. pubescens showed no development. Although Curtis (1942) was able to produce at least protocorms on artificial media, this was not the case though a wide variety of media were used. At the end of the experiment, all seeds appeared to still have live embryos, so non-viable seed is probably not the cause. Approximately 5000 seeds of Pogonia ophioglossoides were planted. One seedling produced a root and a few rudimentary leaflets, and two others produced protocorms by the end of the experiment. This is in contrast to Curtis (1936) and Stoutamire (1964) who described this species as easy to germinate. The results for Calopogon tuberosus were not impressive at the 180 day reading. While Curtis reports obtaining seedlings with leaves 7 cm long after 6 weeks (Curtis, 1936), the longest leaves obtained in this study were about 2 to 3 cm long after the 180 day reading. However, about 50% of the seeds did show some early stages of germination. Spiranthes cernua gave somewhat similar results. A large proportion of the seeds germinated to a small extent. After 180 days, only a few seedlings were vigorous enough to allow removal from the culture tubes. This species and the previous one showed a general decline in vigor after 180 days. The pH of each of the media was checked after the plants were removed from the culture tubes. In every case, the pH had remained constant, thus indicating that the media are sufficiently buffered and that the pH was not a variable factor in the results. The effect of the time period after planting was significant. In general, one would expect good development to continue once started. This is generally what happens in culture of Cattleya and other easily grown genera (Arditti, 1966). Unfortunately, the decline of the vigor of the seedlings in this experiment is not reflected in the growth index readings. The first reading consisted, mainly of green protocorms in Spiranthes, although many of the Calopogon began to develop and already had leaves and roots. The 120 day reading revealed many chlorotic seedlings while the 180 day reading showed a still greater decline, especially in the case of Calopogon. Table 2 shows the analysis of variance for the nutrient medium experiment. Table 3 shows the treatment means for 5 replications and species and medium means. Due to the lack of germination of the Cypripedium species, they were not included in the analysis. Due to the loss of some tubes to contamination, the experiment had to be analyzed with a non-orthogonal program. The species effect was significant at the .01 level for all but the 240 day reading. The


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medium and species x medium effects were significant at the .01 level for all readings. Wynd‘s formula gave the best growth index readings. The reading for the 180 day period was 309. Readings increased with time, indicating a continuous development of the seedlings; however, only one of the seedlings on this medium developed leaves and a root. Many of the seedlings which developed without chlorophyll became necrotic at the time of the last reading. It is impossible to determine exactly when a seedling is dead, so the last reading may be excessively high due to the inclusion of many seedlings that were on the brink of death. The Tsuchiya medium produced the next highest 180 day reading for Spiranthes cernua with a mean of 279. Again, seedlings increased with each reading, showing continuous development. In this case, however, not a single seedling reached stage 6 which can be considered transplanting stage. Many were very chlorotic and appeared as though they would not live much longer. Curtis‘ 1949 medium was third best as rated by the growth index of 256. Again, readings increased for each time period, indicating continued development, but none of the seedlings reached stage 6. All were small, undifferentiated protocorms. Burgeff‘s Eg-1 medium gave a reading of 252 after 180 days. Readings did not change much in the course of the experiment, but the greatest percentage of good seedlings were produced, 2.98% of the seeds produced transplanting-size seedlings (stage 6). Mariat‘s 1952 medium and Liddell‘s medium both gave final readings of 249. In both cases there were many dead seedlings by the end of the experiment. The percent of seedlings at stage 6 were .30% and 0.0% respectively. Kano‘s medium gave a final reading of 248. Many seedlings enlarged to a size greater than those on any other medium. Strong roots and root hairs pushed the seedlings and agar away from the lower side of the culture tubes. The growth index was much lower than would be expected because of the death of many of the seedlings which ceased development at an early stage. Knudson‘s C medium with the addition of 1 mg of niacin per liter of medium gave a reading of 242 after 180 days. Readings were fairly consistent, showing not much net development from period to period. Plants reaching stage 6 was .33% Knudson‘s B medium and the Curtis 1936 medium gave readings of 235 for the final period. The readings were fairly consistent, so there was not much development after the first period. Many seedlings were near death at the end of the experiment. The percent of seedlings at stage 6 was 1.44% and .43 % respectively.


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Table 2

Analysis of Variance 60 Days

__________________________________________________________________

Source d.f. M.S. ―F‖ Blocks 4 67.52 1.27 Species 2 24235.18 455.54 ** Error (a) 8 53.20 Media 19 363.49 15.51 ** Species x Media 38 346.16 14.77 ** Error (b) 184

120 Days

________________________________________________________________

Source df M.S. ―F‖ Blocks 4 204.32 2.37 Species 2 30955.81 359.62 ** Error (a) 8 86.08 Media 19 589.82 4.50 ** Species x Media 38 604.08 4.61 ** Error (b) 184 131.02

180 Days

__________________________________________________________________ Source d.f. M.S. ―F‖ Blocks 4 78.67 2.21 Species 2 37973.36 1066.06 Error (a) 8 35.62 Media 19 866.75 12.03 ** Species x Media 38 816.78 11.08 ** Error (b) 184 73.72

240 Days

__________________________________________________________________ Source d.f. M.S. ―F‖ Blocks 4 298.85 1.11 Species 1 2554.26 9.51 * Error (a) 4 268.59 Media 19 1826.59 6.20 ** Species x Media 19 1333.72 4.53 ** Error (b) 160 294.47

* significant at 5% level ** significant at 1% level


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Table 3 Treatment Means for 5 Replications and Species and Medium Means 60 days 120 days _______________________________ _______________________________ Species Species 1* 2 3 means 1 2 3 means 1 248.0 209.7 200.0 219.2 267.4 206.1 200.0 224.5 2 250.6 199.3 200.0 216.6 266.4 204.2 200.0 223.5 3 211.1 207.3 200.0 216.1 212.4 210.8 200.0 207.7 4 248.0 205.4 200.0 217.8 255.9 211.0 200.0 222.3 5 255.6 207.8 200.0 221.1 258.9 211.0 200.0 223.6 Me 6 234.8 210.1 200.0 214.9 232.8 213.9 200.0 215.6 di 7 218.8 203.7 200.0 207.5 245.2 204.8 200.0 216.6 um 8 214.0 200.5 200.0 204.8 214.0 202.0 200.0 205.3 9 251.6 202.6 200.0 218.1 L.S.D. 273.8 202.7 200.0 225.5 L.S.D. 10 240.8 206.2 200.0 215.7 3.47 236.6 205.6 200.0 214.1 8.19 11 214.2 202.4 200.0 205.5 211.6 204.0 200.0 205.2 12 213.2 206.2 200.0 206.5 214.8 206.6 200.0 207.1 13 228.6 207.2 200.0 211.9 228.2 205.6 200.0 211.3 14 222.4 213.1 200.0 211.8 225.6 216.8 200.0 214.2 15 238.6 206.0 200.0 214.9 238.0 211.4 200.0 216.5 16 254.4 206.3 200.0 220.2 251.2 204.2 200.0 218.2 17 221.6 200.3 200.0 207.3 223.9 202.2 200.0 208.7 18 224.0 200.8 200.0 208.3 224.0 205.6 200.0 209.9 19 223.6 200.0 200.0 207.9 223.6 200.0 200.0 207.9 20 219.4 200.8 200.0 206.7 219.4 200.0 200.0 206.5 Means231.7 204.8 200.0 236.2 206.5 200.0 L.S.D. 2.04 L.S.D 2.59 *1 = Spiranthes cernua, 2 = Calopogon tuberosus, 3 = Pogonia ophioglossoides 180 Days 240 Days ________________________ Species Species 1 2 3 means 2 3 means 1 309.4 205.9 200.1 238.5 244.0 224.7 234.6 2 279.2 199.3 199.1 225.9 224.2 244.0 234.1 3 214.6 205.1 199.9 206.5 237.2 235.6 236.4 4 251.8 207.8 200.0 219.9 232.4 249.5 241.0 5 250.1 201.0 200.0 217.0 218.0 243.8 230.9 6 228.0 210.3 199.9 212.8 230.8 275.5 253.9


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7 225.6 204.0 199.9 209.8 269.0 234.7 251.8 8 219.4 201.7 200.0 207.0 204.7 215.4 210.3 9 227.4 201.6 200.0 209.7 208.1 231.0 219.6 10 241.8 203.0 199.9 214.9 L.S.D 244.0 248.3 246.2 L.S.D 11 233.8 203.6 199.9 214.1 6.15 255.4 216.3 235.8 12.29 12 256.4 206.6 200.0 221.0 237.4 236.2 236.8 13 235.4 202.4 199.9 212.6 259.2 231.2 245.2 14 234.6 205.0 199.9 213.4 302.0 236.6 269.3 15 249.2 211.0 200.0 220.1 239.8 218.0 228.9 16 248.1 205.0 202.0 218.4 223.5 226.0 224.8 17 208.4 199.3 200.1 202.6 217.2 204.3 210.8 18 219.4 198.8 200.0 202.6 219.0 205.5 212.0 19 221.4 197.0 200.0 206.1 207.0 200.0 203.5 20 219.0 199.0 199.9 206.0 257.0 199.7 228.4 Means 238.6 203.4 200.1 236.5 228.4 L.S.D. 176 L.S.D. 4.59

Knudson‘s C medium gave a final reading of 234. This medium produced no seedlings at stage six. A large number of protocorms developed to certain degree and then stopped developing. This was in contrast to a preliminary experiment which showed that this medium produced well-developed seedlings of Spiranthes cernua. Vacin abd Went‘s 1949 medium gave a final reading of 228. Seedlings which developed to stage six in this medium was .96%. The index readings varied little with time. Cosper‘s medium gave a 180 day reading of 227. The readings fell from a high value to a low one because of the death of many seedlings toward the end of the experiment. At first, this medium appeared to be a very promising medium, but apparently something became a limiting factor near the end of the experiment. Seedlings with leaves and roots accounted for 2.29% of the seedlings, although many were chlorotic and appeared to be near death by the end of the experiment. The Thomale medium gave a final reading of 226. Development of the protocorms was poor. Most were colorless and had barely broken through the seed coat when development stopped. No seedlings with leaves and roots were produced. The Vacin and Went tomato juice formulation gave a final reading of 221. The reading in this case may have been quite inaccurate because of the small size of the Spiranthes cernua seeds. The seeds blended in with the tomato fragments in the juice, and were difficult to see. The protocorms were small and appeared bleached.


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The Meyer formulation and the Sladden medium modified by Burgeff gave a final reading of 219. The Meyer tomato juice medium gave results very similar to those described for the Vacin and Went tomato juice medium. The Burgeff medium produced one good seedling, but it was unlike those on other media in that it had very short root hairs and roots. Most of the other protocorms were very small and were beginning to die by the end of the experiment. Burgeff‘s N3f medium gave a 180 day reading of 213. One of the reasons for this low reading is the semi-liquid nature of the medium which made reading the tubes very difficult. No seedlings developed to stage six. The protocorms which did develop were very small and were beginning to die. The germination of Calopogon tuberosus and Pogonia ophioglossoides requires a cold period. About 10 days after the seeds were planted, the embryos swelled noticeably and turned greener. Before the embryos broke through the seed coat, growth ceased in most cases. After 30 days, only a small percentage, usually less than 1% for Calopogon continued to develop and produce leaves and roots. This appears to be a case of double dormancy as described by Barton and Schroeder (1942). In order to overcome this dormancy and get some readings for these species, the culture tubes were removed from the culture chamber and given the cold period described earlier. The results for Calopogon tuberosus were as follows. The 1936 Curtis medium gave the highest growth index reading of 302 after 240 days. The percentage of seedlings which reached stage 6 was 17.40%. This was the highest percentage obtained on any of the media. The Thomale 1954 medium had the second highest percentage of well developed seedlings (those at stage six) 11.35%. In this and the previous case, the growth index was a good indication of which medium produced the most useful seedlings. Knudson‘s B medium gave a growth index reading of 260. The percentage of seedlings at stage 6 was 9.68%. The Meyer tomato juice medium gave the relatively high growth index reading of 254 for Calopogon tuberosus. This reading is highly misleading as is shown by the percentage of seedlings at stage 6, which is 1.18%. As in the case of the other undefined media, the seeds were difficult to see and only large embryos were easy to locate, and this would tend to give a higher growth index reading. Knudson‘s C medium gave a final reading after 240 days of 253. The percentage of seedlings at stage six was 10.88%. Knudson‘s C medium plus 1 mg per liter of niacin gave a final reading of 244 and a percentage of seedlings at stage six of 7.03%. This shows that niacin was not effective in stimulating the germination of this species.


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The Liddell medium gave a growth index reading of 240 after 240 days. The percentage of seedlings at stage six was 11.15% The 1949 Curtis medium was eighth in rank with a growth index reading of 237. None of the seeds reached stage six. Burfgeff‘s N3f medium gave a final growth index reading of 233 for Calopogon tuberosus. This corresponded to 2.82% well developed seedlings at stage six. The Vacin and Went medium gave a final reading of 232 as did the Burgeff Eg-1 medium. This corresponded to a percentage of seedlings at stage six of 4.28% and 5.24% respectively. Wynd‘s medium was in twelfth place with a growth index reading of 229. The percentage of seedlings at stage six was .65%. The seedlings of Calopogon tuberosus on this medium were stunted and most were chlorotic by the end of the experiment. Cosper‘s medium gave a growth index reading of 226. Stage six seedlings represented 6.25% of the total. The Tsuchiya and the Kano media both gave the final growth index reading of 224. This corresponds to .74% and 6.25% of the seed which developed to stage 6. The Calopogon tuberosus seedlings on the Tsuchiya medium were very chlorotic at the end of the experiment, while those on the Kano medium were the best looking seedlings produced on any medium. This is an example of how the growth index values do not necessarily indicate the best seedlings. The Ito medium gave a final reading of 219, and .45% of the seeds developed to stage 6. This sugarless medium seems completely inappropriate for the germination of orchid seed because it does not supply the seeds with a source of carbon. Mariat‘s 1952 medium gave a final growth index reading of 218. None of the seedlings developed to stage six, and all were very chlorotic. The Chang medium gave a final growth index reading of 217, with 2.72% of the seedlings reaching stage six. The percentage of well developed seedlings is undoubtedly high due to the difficulty of seeing ungerminated seeds. The low growth index reading is due to the poor development of the seedlings. The protocorms were extremely chlorotic at the end of the experiment. The Sladden medium modified by Burgeff gave a final growth index reading of 211, and 1.59% of the seeds developed to stage 6. The Vacin and Went tomato juice medium gave the poorest results for Calopogon tuberosus. The growth index reading was 207 with.33% of the seedlings at stage 6. On this undefined medium, the seedlings became chlorotic, and were only slightly developed. The results for Pogonia ophioglossoides are discussed next. The Vacin and Went 1949 medium gave the highest reading after 240 days. The growth index


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reading of 276 corresponded to .319 % of the seedlings which reached stage six. This was the highest percentage for Pogonia ophioglossoides. The reason for the low number appears to be that the 60 days of growth after the cold treatment was not sufficient. At a later time, most of the germinated seeds developed extensive rhizome systems. The Burgeff Eg-1 medium gave a growth index reading of 250. This corresponded to.30% of the seedlings at stage six. Knudson‘s C medium gave the third highest growth index reading of 248, but none of the seedlings developed leaves and roots. Tsuchiya‘s medium number 1 had a final growth index reading of 244. None of the developed seedlings were at stage six, all were at stage three and were developing well. Mariat‘s 1952 medium gave a reading slightly under 244, but none of the seedlings reached stage six. The Curtis 1949 medium number 5 and the Burgeff N3f both gave final growth index readings of 236, and no seedlings developed to stage six on either medium. The Thomale medium gave a final growth index reading of 235, but gave a very high percentage of seedlings at stage six. Both Knudson‘s B and the Cosper medium gave final growth index readings of 231. In both cases, no seeds on either medium reached stage six. The Kano medium gave a final growth index reading of 226. This corresponded to a percentage of well developed seedlings at .176%. Once again, the growth index value does not indicate the development which occurred later. After the last reading, seedlings on this medium almost filled the agar with rhizome systems characteristic of Pogonia ophioglossoides. Wynd‘s medium gave a growth index reading of 225 and there were no well developed seedlings. This is in marked contrast to the effects of this medium on Spiranthes cernua where it gave the highest growth index reading. Liddell‘s medium gave a final growth index reading of 218, with no seedlings reaching stage six. Knudson‘s C medium gave a final growth index reading of 216 with no seedlings at stage six. This is in contrast to the same medium to which 1 mg per liter of niacin was added. The growth index of the medium with niacin was much higher than the growth index reading of the medium without niacin, suggesting that for some orchids, such as Pogonia ophioglossoides, the addition of 1 mg per liter of niacin may increase the germination percentage. The Sladden medium modified by Burgeff gave a reading of 215. No seedlings reached stage six. The Ito medium gave a final reading of 205, with no seedlings reaching stage six. Again, in the case of Pogonia ophioglossoides, we find that the undefined


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media gave poor results. The seedlings were chlorotic and the seedlings which did grow developed only slightly. Chang‘s medium gave a final growth index reading of 204. The seedlings were chlorotic, and none of the seedlings reached stage six. The Vacin and Went tomato juice medium gave a final growth index reading of 200. The same reading applies to Meyer‘s tomato juice medium. No seedling development could be detected.

Discussion and Conclusions This discussion involves the results of Spiranthes cernua after 180 days, and Pogonia ophioglossoides and Calopogon tuberosus after 240 days of growth. The first observation of importance is that the growth index developed by Curtis is not always a very practical method of assessing the results of a test involving these species. The object of this research was to develop a quick method of germinating the seeds of some of our native North American terrestrial orchids which could be used by amateurs or professionals to restock natural areas with native orchid species, and thus preserve the species from extinction. A germination method could also be used by nurseries to propagate orchids for sale to the public. The preservation of rare ecotypes of orchid species would also be possible. Though the growth index is useful for easily grown genera, such as Cattleya, and does give an indication of the developmental progress of these species, it is not of much value in determining which medium might produce the best seedlings, this at stage six. As noted in the results section, frequently many seeds germinated and broke through the seed coat, but then stopped developing. A culture tube in which this had taken place would give a relatively high reading. Another tube in which only a few seeds had germinated and produced leaves and roots, which is necessary for successful culture out of the sterile tubes, would give a relatively low reading, even though it contained useful seedlings while the former tube contained no seedlings of a transplantable size. A more appropriate method for taking readings on these seedlings would simply involve a computation of the percent seeds which produced seedlings of a useful size, those which produce leaves and at least one root. An examination of the media on which Spiranthes cernua produced more than 1% seedlings at stage six shows that they all contained sucrose as the carbon source. This is not true for Calopogon tuberosus and Pogonia ophioglossoides. The greatest percentage of seedling at stage six for Calopogon tuberosus was on the 1936 Curtis medium which was developed for native orchids. It contained 10 gm of glucose as the carbon source. Pogonia ophioglossoides developed best on the Thomale 1954 medium which contained 10 gm of glucose and 10 gm of sucrose. Previous research reviewed by Arditti (1967) shows that for most


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genera, sucrose seemed to be the best sugar for promoting germination, and Kano found that sucrose at 2 to 4% concentrations gave the best results on the genera tested. This appears to be true with Spiranthes cernua, Pogonia ophioglossoides, and Calopogon tuberosus if size of individual seedlings is used as a measure because the largest seedlings were produced on the Kano medium. The results for Spiranthes cernua on Knudson‘s C medium, which contains 20 gm of sucrose per liter are difficult to interpret. A preliminary experiment showed that Spiranthes cernua developed quite well on this medium. The explanation may lie in a physiological change of the seeds during the interval between experiments. In general the media with a concentration of 20 gm of sucrose per liter or more showed good germination. The mineral constituents of the media are quite varied, ranging from completely defined to completely undefined. In some cases apparently dissimilar media gave similar results while similar media gave widely varying results. The media which gave the best results had high levels of nitrate nitrogen, ranging from 16.37 millimoles per liter for Kano‘s medium to 7.03 millimoles for Curtis‘ 1936 medium (see Table 2). However, other media, notably Knudson‘s C and Burgeff N3f had equally high nitrate levels of 12.2 millimoles per liter. These media failed to produce any seedlings at stage six for Spiranthes cernua. Levels of ammonium nitrogen varied from 10.6 millimoles per liter to no ammonium nitrogen. The effect of the ammonium ion is not clear because the results are not consistent. The Cosper medium lacked this ion but produced the highest growth index, and the third highest percentage of seedlings at stage six for Spiranthes cernua. However the seedlings lacked vigor and were very chlorotic. This may be due to a lack of ammonium nitrogen, but according to Raghavan and Torrie (1964), this ion is only beneficial in the early seedling stages of Cattleya, and once the early stage is passed, it seems to be of no benefit. The level of potassium varied from .882 to 9.16 millimoles per liter. The best development as measured by the percent seedlings reaching stage six, was at the higher levels of potassium, but again the results were not consistent. The level of the phosphate ion varied from .112 to 3.27 millimoles per liter. Again, the best development was associated with high levels of this ion. The Cosper medium had the lowest level of this ion, and this may have contributed to the general lack of vigor of the seedlings on this medium. Although it has been suggested that a high level of the phosphate ion might cause the precipitation of iron to the point that growth is inhibited (Arditti, 1967) this appears not to be the case in this experiment. The level of iron in the media varied from zero to .89 millimoles per liter. The Kano medium contained no additional iron, but presumably the impure


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nature of the chemicals which make up this commercial fertilizer contain sufficient iron because seedlings on this medium showed no sign of chlorosis. The level of sulphate and calcium ions seems to bear no relation to the development of the seedlings, however most of the media which produced more than 1% good Spiranthes cernua. Seedlings contained added magnesium. The Kano medium was the exception again. Magnesium may have been present in the impure chemicals in adequate supply. Chlorine and manganese had no apparent effects. In summary, the nutritional requirements for a medium to produce vigorous seedlings of Spiranthes cernua, Calopogon tuberosus, and Pogonia ophioglossoides probably include sucrose at 20 to 35 gm per liter, nitrate nitrogen at a concentration of 10 to 16 millimoles per liter, ammonium at 3 to 7.5 millimoles per liter, phosphate at 1.2 to 3.2 millimoles per liter, iron at .2 to .9 millimoles per liter, calcium at 1.4 to 3.2 millimoles per liter, and magnesium at 1 to 1.5 millimoles per liter. Presumably a medium with these characteristics should allow good germination and development of these species, although much variability is evident in these species, and the exact conditions which trigger the development of seedlings with leaves and roots not completely clear. The storage time of the seeds probably had some effect on germination due to a biological clock which governs the time from capsule dehiscence to germination.

Acknowledgements

The author wishes to thank Dr. Ervin Denisen for his efforts to secure an NDEA Title IV teacher training fellowship which allowed the author to perform this research. Dr. Charles Sherwood provided a review of this paper. Dr. Jowett set up the statistical experimental designs, and analyzed the data. Dr. L.Y. Quinn supplied some of the seeds used in these experiments. Thanks also to Chuck Sheviak for thought-provoking discussions of what might be the causes of poor germination in our native North American terrestrial orchids.

Literature Cited Arditti, J. 1966. The effects of niacin, adenine, ribose and niacinamide coenzymes on

germinating orchid seeds and young seedlings. Amer. Orch. Soc. Bul. 35: 892-898. Arditti, J. 1967. Factors affecting the germination of orchid seeds. Bot. Rev. 33: 1-97. Barton, L. V. and E. M. Schroder. 1942. Dormancy of seeds of Convallaria majalis L. and

Smilacina racemosa (L.) Desf. Contrib. Boyce Thompson Institute 6: 277-230. Burgeff, H. 1936. Die Samenkeimung der Orchideen. G. Fischer, Jena. Curtis, J. T. 1936. The germination of native orchid seeds. Amer. Orch Soc. Bul. 5: 42-47. Curtis, J. T. 1937. Non specificity of orchid mycorrhizal fungi. Proc. Soc. Exp. Biol. and Med.

36: 43-44. Curtis, J. T. 1939. The relation of specificity of orchid mycorrhizal fungi to the problem of

symbiosis. Amer. Jour. Bot. 26: 390-399.


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Curtis, J. T. 1942. Germination and seedling development in five species of Cypripedium L. Amer. Jour. Bot. 30: 199-205.

Fisher, R. A. and P. Yates. 1953. Statistical Tables. Oliver and Boyd. Edinburgh. Fernald, M. L. 1950. Gray’s manual of botany. American Book Co. New York. Galston, A. W. 1961. The life of the green plant. Prentice Hall, Inc. New York. Israel, H. W. 1963. The production of Dendrobium seedlings by asceptic culture of excised

ovules. Amer. Orch. Soc. Bul. 32: 441-443. Kano, K. 1968. Acceleration of germination of the so-called ―hard to germinate‖ orchid

seeds. Amer. Orch. Soc. Bul. 37: 690-698. Knudson, L. 1922. Non symbiotic germination of orchid seeds. Bot. Gaz. 73: 1-25. Knudson, L. 1924. Further observations on nonsymbiotic germination of orchid seeds. Bot.

Gaz. 77: 212-219. Knudson, L. 1925. Physiological study of the symbiotic germination of orchid seeds. Bot.

Gaz. 79: 345-379. Knudson, L. 1930. Flower production by orchid grown non-symbiotically. Bot. Gaz. 89: 192-

199. Knudson, L. 1941. Germination of seed of Goodyera pubescens. Amer. Orch. Soc. Bul. 9: 199-

201. Laetsch, W. M. and R. E. Cleland. 1967. Papers on plant growth and development. Little, Brown

and Co, Boston. Liddell, R. W. 1944. Germinating native orchid seed. Amer. Orch. Soc. Bul.: 12. 344-345. Miller, C. O. 1956. Similarity of some kinetin and red light effects. Plant Physiol. 31: 318-319. Niemann, D.A. 2001. Orchid Propagation. Orchids 68: 460-470. Noggle, C. R. and F. L. Wynd. 1943. Effects of vitamins on germination and growth of

orchids. Bot. Gaz. 104: 455-459. Northen, R. T. 1962. Home orchid growing., 2nd ed. Van Nostrand, Princeton. Raghavan, V. and J. G. Torrey. 1964. Inorganic nitrogen nutrition of the seedlings of the

orchid Cattleya. Amer. Jour. Bot. 51: 264-274. Redlinger, J. R. 1961. Sterilizing agents for orchid seed flasking. Amer. Orch Soc. Bul. 30: 800-

801. Snedicor, G. W. and W. G. Cochran. 1967. Statistical methods. 6th ed. Iowa State University

Press, Ames. Spoerl, E. 1948. Amino acids as sources of nitrogen for orchid embryos. Amer. Jour. Bot. 35:

88-95. Withner, C. L. 1959. The orchids. The Ronald Press. New York. Stoutamire, W. P. 1964. Seeds and seedlings of native orchids. The Michigan Botanist. 3: 107-

119.


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Appendix 1.

Composition of the Media Burgeff EG-1

Dilute solution A and B in 500 ml water, then combine solutions Solution A

Calcium nitrate Ca(NO3)2.4H20 ...................................... 1.00 gm Ammonium sulphate (NH 4)2SO4 ................................... .25 Magnesium sulphate MgSO4.7H2O................................... .25

Solution B Ferrous sulphate FeSO4.7H2O ........................................... .02 Potassium phosphate KH2PO4 ........................................ .25 Potassium phosphate K2HPO4 .......................................... .25 Sucrose ............................................................................... 20.00 Agar (Bacto) ....................................................................... 15.00

Burgeff N3f Part A

Magnesium sulphate MgSO4.7H2O................................... .25 gm Potassium chloride KCl ........................................................ .25 Ferrous sulphate FeSO4.7H2O ....................................... .02 Calcium nitrate Ca(NO3)2.4H2O .................................... 1.00 Ammonium sulphate (NH4)2SO4 .................................... .25

Part B Citric acid .............................................................................. .09 Potassium phosphate K2HPO4.3H20 ............................... .25

Dissolve parts A and B in 500 ml water, combine solutions, then add: Glucose ............................................................................... 10.00 Fructose .............................................................................. 10.00 Agar (Bacto) ....................................................................... 12.00

Adjust pH to 5.0

Chang Fish emulsion* ..................................................................... 1.5 teaspoons Agar (Bacto) ......................................................................... 9.0 Sugar (Sucrose was used ) ................................................ 5.5 Peptone (Bacto) ................................................................ 1.0 Water .............................................................................. 1000 ml

* Atlas is recommended, but Ortho was substituted; the analysis is 5-2-2

Cosper Potassium nitrate KNO3 ................................................ .820 gm Calcium nitrate Ca(NO3)2 ......................................... .236 Potassium phosphate KH2PO4 .................................. .024 Potassium chloride KCl ............................................... .065 Magnesium sulphate MgSO4.7H2) ............................ .036 Ferric tartrate ..................................................................... .0015 Urea .................................................................................... .050 L asparagines ..................................................................... .050 Sucrose ........................................................................ 20.00 Agar (Bacto) ............................................................. 15.00 Water ............................................................................ 1000ml

pH adjusted to 5.0


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Curtis 1936

Monopotassium phosphate KH2PO4 ...................... .12 gm Magnesium sulphate MgSO4.7H2O ........................... .26 Ammonium nitrate NH4NO3 .................................. .22 Calcium nitrate Ca(NO3)2.4H2O ............................... .35 Ferric phosphate FePO4 .............................................. .003 Glucose ............................................................................ 10.00 Agar (Bacto) ................................................................ 14.00 Water .......................................................................... 1000 ml

pH adjusted to 4.7

Curtis 1949 Potassium phosphate KH2PO4 ................................. .112 gm Magnesium sulphate MgSO4.7H2O ........................ .260 Calcium nitrate Ca(NO3)2.4H2O ............................ .350 Ammonium nitrate NH4NO3 .................................... .220 Ferric phosphate FePO4 ................................................. .027 Yeast extract (Bacto) ........................................................ .10 Agar (Bacto) ............................................................ 15.00 Water ............................................................................ 1000 ml

pH adjusted to 4.8

Ito Corn starch (Argo) ......................................................... 50.0 gm Libby‘s tomato juice .....................................................100.0 ml

Add water to make 500 ml

Kano Hyponex ......................................................................... 3.0 gm Pepone (Bacto) ............................................................ 2.0 Sucrose .......................................................................... 35.0 Agar (Bacto) ............................................................... 15.0 Water ............................................................................ 1000 ml

pH adjusted to 5.0

Knudson‘s B Potassium phosphate KH2PO4 ................................. .25 gm Calcium nitrate Ca(NO3)2 ........................................... 1.00 Ammonium sulphate (NH4)2SO4 .......................... .50 Magnesium sulphate MgSO4.7H2O ....................... .25 Ferric phosphate FePO4 ............................................ .05 Sucrose .......................................................................... 20.00 Agar (Bacto) ............................................................... 17.5 Water ........................................................................... 1000 ml

pH adjusted to 5.0

Mariat 1952 Magnesium sulphate MgSO4.7H2O .......................... .25 gm Calcium nitrate Ca(NO3)2.4H2O ........................... 1.00 Monopotassium acid phosphate KH2PO4 ............ .25 Calcium chloride CaCl2.2H2O ................................. .12 Ferric chloride FeCl3 ................................................. .001 Agar (Bacto) ............................................................... 16.00 Sucrose ........................................................................ 20.00 Water ........................................................................... 1000 ml

pH 5.0


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Meyer 1945

Fresh strained tomato juice ..................................... 250 ml Agar (Bacto) ................................................................ 7.5 gm Water ......................................................................... 250 ml

pH 4.8

Sladden modified by Burgeff Calcium nitrate Ca(NO3)2.4H2O ................................ 1.00 gm Magnesium sulphate MgSO4.7H2O .......................... .30 Monopotassium acid phosphate KH2PO4 ........... .40 Ferric citrate .................................................................... .20 Ammonium sulphate (NH4)2SO4 .......................... .70 Potassium citrate ........................................................... .35 Niacin ............................................................................. .001 Fructose ........................................................................ 10.00 Glucose ......................................................................... 10.00 Agar (Bacto) .............................................................. 12.00 Water .................................................................. 1000 ml

pH 5.0

Thomale 1954 Ammonium sulphate (NH4)2SO4 ....................... .06 gm Ammonium nitrate NH4NO3 ............................... .37 Potassium nitrate KNO3 ...................................... .40 Potassium phosphate KH2PO4 .......................... .30 Magnesium nitrate Mg(NO3)2.6H2O ............... .11 Ferrous sulphate FeSO4 ...................................... .02 Fructose .................................................................... 10.00 Glucose ..................................................................... 10.00 Agar (Bacto) ............................................................ 16.00

pH 5.0

Tsuchiya 1954 Magnesium sulphate MgSO4.7H2O ...................... .36 gm Calcium nitrate Ca(NO3)2.4H2O ......................... .20 Potassium nitrate KNO3 ......................................... .08 Sodium sulphate Na2SO4 ........................................ .20 Potassium chloride KCl ............................................ .065 Potassium phosphate KH2PO4 .............................. .0165 Manganese sulphate MnSO4 .................................... .0045 Zinc sulphate ZnSO4.7H2O ................................... .0015 Boric Acid H3BO3 ...................................................... .0015 Potassium iodide KI .................................................. .00075 Glycine ............................................................................. .003 Nicotinic acid ................................................................ .0005 Pyridoxine ..................................................................... .0001 Thiamin ........................................................................ .001 Sucrose ............................................................................ 20.000 Agar (Bacto) ............................................................... 15.00 Water ........................................................................... 1000 ml

pH 5.5


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Vacin and Went 1949

Tricalcium phosphate Ca3(PO4)2 ................................... .20 gm Potassium nitrate KNO3 ..................................................... .525 Monopotassium acid phosphate KH2PO4..................... .25 Magnesium sulphate MgSO4.7H2O ................................ .25 Ammonium sulphate (NH4)2SO4 ................................... .50 Ferric tartrate .......................................................................... .028 Magnesium sulphate MgSO4.7H2O ................................ .0075 Sucrose ................................................................................. 20.00 Agar (Bacto) ....................................................................... 16.00

pH 5.0

Vacin and Went tomato formulation Fresh tomato juice .......................................................... 375 ml Sucrose .................................................................................. 5.0 gm Agar (Bacto) ......................................................................... 7.5 Water .................................................................................. 125 ml

pH 4.8

Wynd 1933 Type iv R5S1 Potassium sulphate K2SO4 ......................................... .435 gm Monocalcium acid phosphate CaHPO4 .................... .327 Magnesium nitrate Mg(NO3)2.6H2O ..................... .998 Manganese sulphate MnSO4 ..................................... .000669 Ferric phosphate ............................................................ .000269 Sodium borate Na2B4O7.10H2O ........................... .00143 Maltose ......................................................................... 20.00 Agar (Bacto) ............................................................... 17.5 Water .......................................................................... 1000 ml

pH 4.9

Knudson‘s C Potassium phosphate KH2PO4 ..................................... .25 gm Calcium nitrate Ca(NO3)2.4H2O................................... 1.00 Ammonium sulphate (NH4)2SO4 .................................. .50 Magnesium sulphate MgSO4.7H2O ............................... .25 Ferrous sulphate FeSO4.7H2O ....................................... .025 Manganous sulphate MnSO4 ........................................... .0075 Agar (Bacto) ....................................................................... 17.5 Water .............................................................................. 1000 ml

pH adjusted to 5.0 (For Knudson‘s C plus niacin added, 1 mg of niacin was added per liter of the above solution)

Liddell

Calcium nitrate Ca(NO3)2.4H2O ........................... 1.00 gm Ammonium nitrate NH4NO3 ................................ .25 Magnesium nitrate MgSO4.7H2O .......................... .25 Potassium phosphate KH2PO4 .............................. .20 Potassium phosphate K2HPO4 .............................. .10 Glucose .................................................................. 18.00 Sucrose ................................................................... 2.00 Agar (Bacto) .......................................................... 15.00 Water .................................................................. 1000 ml

pH adjusted to 5.0

Niemann: II. EFFECTS OF VARIOUS GROWTH REGULATORS ON THE GERMINATION

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II. The Effects of Various Growth Regulators on the Germination and Development of Several North American

Native Orchids

David Niemann

Introduction The North American native terrestrial orchids face threats from agriculture and urbanization. Many habitats have been destroyed resulting in the extinction of local ecotypes of many of the species. Setting aside areas where these species can continue to grow would be the best way to preserve the species and ecotypes, but few individuals or organizations have the financial ability to do this. As a second choice, these orchids could be grown from seed and kept in cultivation until suitable areas are set aside where they can be planted and be allowed to grow Niemann, 2001. Some species of North American orchids do not germinate well, possibly because of dormancy problems. This paper reports the results of the use of several growth regulators in an attempt to break dormancy of these seeds.

Materials and Methods The seed of Cypripedium reginae were collected in my garden in Elmwood Park, Illinois on September 1, 1968. The capsules were still green at the time of harvest. Seeds of Calopogon tuberosus were collected in Lake County, Illinois on September 20, 1968. Seeds of Cypripedium acaule were donated by Dr. L. Y. Quinn, of Iowa State University. Seeds were removed from the capsules, air dried, and placed in a closed jar containing calcium chloride. The jars were kept in a refrigerator at 4 degrees C until the seeds were planted. The seeds were sterilized in a 2%Clorox (5.25 % sodium hypochorite) according to Redlinger (1961). The seeds were shaken vigorously for 10 minutes in a 1:50 solution of clorox:water. The seeds were then rinsed twice in sterile distilled water. The seeds these species could not be suspended in water. They were planted by removing about 100 seeds from the sterilization vial with a small spatula. The seeds were then injected into a culture tube with a stream of sterile distilled water. With some practice, fairly uniform quantities of seed could be sown in each tube.


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The tubes were placed in a growth chamber with approximately 200 foot-candles of fluorescent light, and a temperature of approximately 24 degrees C. Three readings were taken on each tube at 60 day intervals after planting. The readings were taken using the growth index developed by Curtis (described by Spoerl, 1948). In previous orchid research results were difficult to interpret because a uniform method of taking data was not used. Knudson and other early workers used measurements such as fresh and dry weight of seedlings, seedling color, protocorm diameter, total leaf and total root length. These characteristics may be all right for some purposes, but only one reading can be taken on a group of seedlings. The growth index is probably the best method of taking data because several readings can be taken on the same group of seedlings without destroying them. A major advantage is distinguishing degrees of development. Protocorm diameter is misleading because a large protocorm is as undeveloped as a small one. The growth index takes into account the developmental stage of a seedling. There are six stages in my modification of the index. These may be seen in Figure 1. The index is calculated by counting the number of seedlings at each stage in a culture tube. From this the percent of seedlings at each stage is calculated. Then the percent of seedlings at each stage is multiplied by the stage number. The sum of these products is the growth index for that tube. The data obtained in this manner are suitable for statistical analysis (Spoerl, 1948). The basal medium used in this experiment was White‘s standard with 15 gm agar and 20 gm of sucrose per liter of medium (see appendix). The growth regulating substances were prepared by dissolving the proper amount of the chemical in 95% ethyl alcohol to make a one thousandth molar solution. This was then added to the nutrient media at a temperature of 45 degrees C. to give a final concentration of one millionth molar. Statistical Design The experiment was set up in a factorial design. There were two blocks of two species. The treatments were gibberellic acid (GA-3); kinetin; 2,4-D; and indole-3-acetic acid. They were present alone and in all possible combinations. This gave a total of 16 treatment combinations. The readings were taken using the growth index. Results The seeds of Cypripedium reginae and C. acaule did not germinate on any of the media. Calopogon tuberosus germinated on most media, and gave a general idea of what the effects of these growth regulators are. Statistical analysis was not possible because of the small number of seeds which actually germinated. Most of the culture tubes produced only 4 or 5 seedlings, so interpretation must be tentative until a means to produce more seedlings is developed.


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The check culture tubes, those with no growth regulators added, produced seedlings that were abnormal. One tube contained a mass of callus about 1 cm in diameter. This type of development was not noted in the culture tubes of any previous experiments. Several small protocorms were produced, but none developed leaves or roots. Gibberellic acid alone produced very large protocorms, up to 3 mm in diameter. Some seedlings produced leaves. In at least one case, a seedling went dormant as described by Stoutamire (1964). It then resumed growth without the requirement of a cool period. This had not been observed in any previous experiments described in Niemann (In press). The kinetin medium produced one plant with a root, but rhizomes were very short. The leaves were about one third normal size, although some large corms up to 3 mm in diameter developed. Kinetin with gibberellic acid also gave some evidence of a bypass of the dormant period requirement, but the effect was not as clear as that mentioned for gibberellic acid alone. Some large corms 3 mm in diameter developed. The medium with 2,4-D alone caused very short rhizoids less than 1 mm long developed on Calopogon tuberosus. There was an apparent disorganization of the shoot apical meristem. The seedlings were not normal, and some consisted of simple linear tissue masses. Plants on the medium including 2.4-D and gibberellic acid developed very little, only to the protocorm stage, and then most of the protocorms died. The medium with 2,4-D caused the production of one seedling with three stolon-like growths not characteristic of Calopogon tuberosus. The rest of the seedlings showed a disruption of the shoot apical meristem. Many of the seedlings died The medium with indole acetic acid alone caused poor root growth. The leaves were about twice the normal length, and many of the seedlings had died by the end of the experiment. The medium with indole acetic acid plus gibberellic acid caused some of the corms to be very large, up to 4 mm in diameter. There was again some evidence that the dormant period was being overcome, but not so strongly as on the medium with gibberellic acid alone. Leaf and root growth were normal. The medium with indole acetic acid plus kinetin prevented roots from forming. The leaves were very short, only about 20% of the normal length. The leaves formed a rosette over the plant. The medium containing indole acetic acid plus gibberellic acid plus kinetin produced leaves of normal size, but large corms up to 3 mm in diameter. The medium containing indole acetic acid plus 2.4-D caused very poor development. Many of the protocorms died and several linear, undifferentiated seedlings developed.


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The medium containing indole acetic acid plus 2,4-D plus gibberellic acid caused many seedlings to die and others became linear seedlings. Development was not as good as that of the above treatment. The medium with indole acetic acid plus 2,4-D plus kinetin caused the development of many protocorms, but they failed to grow. Many died by the end of the experiment. The medium with indole acetic acid, gibberellic acid, kinetin and 2,4-D caused the development of many protocorms, but most had died by the end of the experiment.

Other Tests A few other tests were performed and they will be reported briefly. One experiment involved soaking seed of Calopogon tuberosus for 45 days in sterile distilled water. Liddell (1944) has suggested that soaking seed of this species for about a month in nutrient solution enhanced germination. Soaking seed in sterile distilled water should remove any water-soluble germination inhibitors which may be present. In this experiment, germination was poorer when the seed was soaked in water. It appears that Calopogon tuberosus does not possess a water-soluble germination inhibitor. In another test, seeds of Cypripedium acaule were sown on media with and without a saponin extracted from Cypripedium acaule by Dr. L. Y. Quinn of Iowa State University. The substance was believed to have some stimulatory effect on the germination of orchid seeds. After 180 days in culture, no development of protocorms was noted. In another experiment, indole –3-acetic acid, gibberelic acid and kinetin were used individually at concentrations of one thousandth, one ten thousandth, one one-hundred thousandth, and one millionth molar. Seeds of Spiranthes cernua, and Cypripedium acaule were planted. The experiment was concluded at the first reading because none of the treatments appeared to give any beneficial effect on inducing seed germination.

Discussion and Conclusions Test of Growth Regulators

General trends were observed which seem to follow from previous work on other species. The discussion will be limited to the main effects because the interactions appeared to be a simple blend of these effects. Statistical analysis was not performed because of the small number of seeds which germinated,

Gibberellic Acid The apparent bypass of the need for a dormant period is in agreement with the observations in other plants. Sachs (Laetsch and Cleland, 1967) states that gibberellic acid can substitute for a cold period to induce bolting of the rosette plant Hyoscyamus niger var. bienne, and states that other workers have made


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similar discoveries for other plants. Thus it appears that of the growth regulators tested, gibberellic acid is the only beneficial one in that it allowed more or less continuous development of Calopogon tuberosus without the interruption of a cold period. It was hoped that gibberellic acid would also stimulate ungerminated seeds to germinate. Kahn (1957) found that gibberellic acid was able to break dormancy of lettuce seeds whose germination was prevented by temperature inhibition, dark-osmotic inhibition, or far red light inhibition. Apparently gibberellic acid at the concentrations used in this experiment is not capable of substituting for the cold period which is required to bring about the germination of Calopogon tuberosus. Since gibberellic acid gave no indication of stimulating the germination of Cypripedium seeds, its buildup in the seeds is probably not required for germination as it is in some seeds, such as Avena (Galston, 1961). Kinetin The effect of kinetin for the most part was inhibitory. Other work has shown that its effect on tropical orchids is also inhibitory. Miller (1956) observed that its stimulation of lettuce seed germination in the light and darkness, however in this experiment, stimulation of orchid seed germination did not occur. 2,4-D The auxin 2,4-D was completely non-beneficial in this study. It apparently induces cell division in Calopogon tuberosus in the concentration used, but cell division is erratic and abnormal. Differentiation did not take place. 2,4-D also seems to have a detrimental effect on the metabolism of this monocot species, indicating that 2,4-D may be a factor in the local extinction of this species and perhaps other orchids. Indole acetic acid This chemical apparently has very little effect on Calopogon tuberosus. In combination with gibberellic acid it may cause slightly larger seedlings to develop. It has been noted that indole acetic acid causes an increase in the fresh weight of the seedlings of some tropical orchids. If enough seeds had germinated in this experiment an increase in fresh weight might also have occurred. The general effect of indole acetic acid in the increase in cell wall plasticity and elasticity in preparation of cell walls before division did not occur in this experiment. In conclusion, the only additive used in this experiment which was beneficial at the one hundred thousandth moles per liter was gibberellic acid. It helps override the requirement for a cool dormant period.


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Acknowledgements The author wishes to thank Dr. Ervin Denisen for his efforts to secure an NDEA Title IV teacher training fellowship which allowed the author to perform this research. Dr. Charles Sherwood provided a review of this paper. Dr. Jowett set up the statistical experimental designs, and analyzed the data. Dr. L. Y. Quinn supplied some of the seeds used in these experiments. Thanks also to Chuck Sheviak for thought-provoking discussions of what might be the causes of poor germination in our native North American terrestrial orchids.

Literature Cited Galston, A. W. 1961. The life of the green plant. Prentice Hall, Inc. New York. Kahn, A. J., A. Goss, and D. E.. Smith.1957. Effect of gibberelic acid on germination of

lettuce seeds. Science 125:441-443. Laetsch, W. M. and R. E. Cleland. 1967. Papers on plant growth and development. Little, Brown

and Co, Boston. Liddell, R. W. 1944. Germinating native orchid seed. Amer. Orch. Soc. Bul.: 12. 344-345. Niemann, D. A. 2001. Orchid Propagation. Orchids 68: 460-470. ------. The effects of various media on the germination and development of several North

American native orchids (in press) Redlinger, J. R. 1961. Sterilizing agents for orchid seed flasking. Amer. Orch Soc. Bul. 30: 800-

801. Spoerl, E. 1948. Amino acids as sources of nitrogen for orchid embryos. Amer. Jour. Bot. 35:

88-95. Appendix 2.

White‘s standard (for culturing plant and animal cells)

Calcium nitrate Ca(NO3)2.4H2O ................................................. 12.0 gm Potassium nitrate KNO3 ................................................................. 3.2 Potassium chloride KCl .................................................................. 2.6 Magnesium sulphate MgSO4.7H2O ............................................ 30.0 Sodium sulphate Na2SO4 ............................................................... 8.0 Sodium phosphate NaH2PO4.H2O ............................................... .76 Manganese sulphate MnSO4.7H2O ................................................ .20 Zinc sulphate ZnSO4.7H2O .............................................................. .12 Boric acid H3BO3 ................................................................................ .06 Potassium iodide KI ............................................................................ .03 Copper sulphate CuSO4 .................................................................... .0004 Molybdic acid MoO3 (85%)............................................................. .00004 This is added to 4000 ml of water and serves as a stock solution which is further diluted by adding 100 ml of stock solution to 899 ml water. The following is dissolved in 100 ml of water and serves as a stock solution: 1 ml is added per liter of final solution. Glycine ..................................................................................................... .300 gm Nicotinic acid........................................................................................... .050 Thiamin hydrochloride .......................................................................... .010 Pyridoxine hydrochloride ...................................................................... .010 Sucrose .................................................................................................. 20.00 Agar (Bacto .......................................................................................... 15.00

Adjust pH to 5.5

David Neimann, 9517 Seeman Road, Union, Illinois 60180 [email protected]

mailto:[email protected]


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Note of Interest:

Epipactis palustris (L.) Crantz another European visitor

new to the North American orchid flora On June 10, 2006 Mark Larocque

found plants in bud of an Epipactis which he suspected would be the European E. palustris growing in a flooded limestone area near the Susquehanna River in Lancaster County, Pennsylvania. When he checked the site a few weeks later the plants has finished flowering and were in fruit. Returning on June 16, 2007 Mark found the plants in bud again. Due to prior commitments he was to be unable to return to the site until July 1. Tom Nelson from New York City visited the site on June 25 and found the plants in flower confirming the identity of the Epipactis. The population appears to be reproducing and the plants are growing in a

protected area along with Liparis loeselii and Spiranthes lucida. This is the first confirmed record for this species for North America.

In Europe Epipactis palustris occurs rather frequently in calcareous marshes and is also popular with native orchid growers. The New York Flora Atlas lists the plant for New York based upon a 1990 collection of a specimen taken from a population in cultivation near Albany that is spreading in the owner‘s yard, although it has yet to appear spontaneously in the vicinity. Weldy, T., R. Mitchell, and R. Ingalls. 2002. New York Flora Atlas. New York Flora Association, New York State Museum, Albany, NY. http://nyflora.org/atlas/atlas.htm Photos by Tom Nelson

Epipactis palustris: another European visitor

http://nyflora.org/atlas/atlas.htm


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TEXAS LADIES’ TRESSES, SPIRANTHES BREVILABRIS, REDISCOVERED IN TEXAS

Eric Keith

On April 11, 2007, while installing vegetation monitoring plots for the Sam Houston National Forest in Walker County, Texas, I noticed a small flowered Spiranthes with persistent oval basal leaves. I knew that this combination of characteristics could be only one of three rare spring flowering species known from Texas. The densely pubescent rachis and yellowish central lip distinguished the species as Texas ladies‘-tresses, Spiranthes brevilabris. The other two species are Florida ladies‘-tresses, Spiranthes floridana, and Eaton‘s ladies‘-tresses (Spiranthes eatonii) (Brown and Folsom 2003; Sheviak and Brown 2002). On two subsequent visits to the site (April 14 and April 17), I recorded 21 additional plants. The habitat for this population is a remnant black clay (blackland) prairie approximately five acres in size. Several blackland prairies are found in Walker County and the Sam Houston National Forest, but no plants were found on any of the other prairies surveyed. This species was originally described from southeastern Texas in 1840 by John Lindley based on a specimen collected by Thomas Drummond (Liggio and Liggio 1999). Historically, the species has been collected in four other Texas counties with the most recent collection from Galveston County in 1975 (Liggio and Liggio 1999, Digital Flora of Texas 2003). Until this rediscovery, no surviving populations were known from Texas, and only two other extant sites are known for the species, both occurring in Florida (Paul Martin Brown, personal communication). The habitat for the species has been described as sandy soil in moist prairies, pine-hardwood forest, and wetland pine savannahs (Liggio and Liggio, 1999) and dry to moist roadsides and fields (Sheviak and Brown, 2002). Since the species is now known to also occur on blackland prairies, it is hoped that further surveys in suitable habitat will yield additional populations. I would like to thank Paul Martin Brown for confirming the identity of the plants. Literature Cited: Brown, P.M. and S.N. Folsom. 2003. The Wild Orchids of North America, North of Mexico.

Gainesville: University Press of Florida. Digital Flora of Texas. 2003. Herbarium Specimen Browser website.

http://www.csdl.tamu.edu/FLORA/tracy2/main1.html Liggio, J and A.O. Liggio. 1999. Wild Orchids of Texas. University of Texas Press: Austin.

Keith: TEXAS LADIES’ TRESSES, SPIRANTHES BREVILABRIS, REDISCOVERED IN TEXAS



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Sheviak, C.J. and P.M. Brown 2002. Spiranthes. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 26, pp. 530-545.

Eric Keith, Raven Environmental Services, Inc., P.O. Box 6482, Huntsville, TX 77342, [email protected]

Blackland prairie, Walker Co., Texas; with the author‘s sons Lance and Will

Spiranthes brevilabris, Walker Co., Texas April 11, 2007



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BOOK REVIEWS Orchids of Europe, North Africa and the Middle East Pierre Delforge 2006 Timber Press Full color hardcover, 640 pp., 5 x 7.5 in (190 x 125 mm); 1270 color photos plus several line drawings and watercolors ISBN-13: 9780881927542; ISBN-10: 0881927546 $39.95 Order from: http://www.timberpress.com/books/isbn.cfm/0-88192-754-6/orchids_europe_north_africa_middle_east/delforge Delforge‘s previous works have always been well received as they should be. He work is meticulous and although limited to brief entries for each species, as complete as space permits. The gallery of photographs is mind-boggling. This current book is no exception. It combines the best of all of his other books and presents a single volume that is both an update of previous works and much new material as well as needed revisions both taxonomically and nomenclaturally. Technically a 3rd edition of his original work of the same name it has more than 200 additional pages than previous French editions. Introductory chapters on basic orchid information—anatomy, life cycles, reproduction and orchid identification are detailed and easy to understand with many excellent graphics. The following chapters of species accounts are broken into four chapters and arranged by subtribes and the genera arranged systematically. Although this arrangement is excellent for the botanist it does make it difficult to find a specific genus or species quickly without consulting the index. Because of the large geographic area covered by the book one must read through the descriptions very carefully to find those species that might occur where the reader is exploring. This is not necessarily a negative aspect of the book but does force the reader into review many more species. A few synonyms are given with each species entry and fortunately they are in the index as well. Several taxonomical and nomenclatural points should be mentioned. The genus Listera is nested into Neottia, Pseudorchis into Gymnadenia; although Coeloglossum is maintained in its own genus rather than in Dactylorhiza. Hybrids are treated extensively as are aberrant and unusual forms such as albinos, white-flowered forms and other various color variants. The largest genera treated—Epipactis, Dactylorhiza, Orchis, and Ophrys are as complete as they can possibly be with extensive illustrations to help sort out the oft-confusing species and forms. Ophrys alone has over 250 pages devoted to the genus! Pierre Delforge has studied orchids, observed the evolution of their habitats, and protected them for more than 35 years in Europe, North Africa, and the Middle East and is an expert on European orchids for the IUCN — the World Conservation Union. I can recommend this new work with no reservations and at $39.95 it is one of the best buys in orchid literature available! PMB NOTE: This title has been published in Great Britain by A & C Black (ISBN 9780713675252) and cooperatively with Timber Press in the North America. They have different cover images but the body of the books is the same.

http://www.timberpress.com/books/isbn/9780881927542/orchids_europe_north_africa_middle_east/delforge


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ORCHIDS IN HAWAII Ted Green $10.95 paper 6" x 9" 130 pp. 175+ color photographs ISBN: 1-56647-720-4 Mutual Publishing, LLC 1215 Center Street, Suite 210, Honolulu, HI 96816

Hawai‗i‘s love affair with orchids began in the early 1950‘s when a few enterprising growers made the delicate plants affordable for your everyday, ―garden variety‖ hobbyist. Today, Hawai‗i is famous for its orchids. With only three of its own native species out of the more than 35,000 species of orchid world wide, it seems unlikely but true. Orchids in Hawai‗i offers an introduction to orchids and explains where and when they can be found, describes species, hybrids, and how to read labels as well as how to take plants to the mainland. Filled with over 175 stunning photos it is truly the best guide a budding orchid enthusiast could pick up.

Orchid Arabia Eric Hansen; illustrated by Barbara Evans Saudi Aramico World November/December 2006 pp. 10-16 http://www.saudiaramcoworld.com/issue/200606/orchid.arabia.htm Having nothing to do with North American orchids, but nevertheless a fascinating and beautifully illustrated article that may be accessed electronically. Hansen is the author of the popular Orchid Fever. ORCHIDELIRIUM Harold Feinstein with an Introduction by Robert H. Hesse Bulfinch Press February 2007 $50.00 144 pages 100 full color photographs 13.5 x 11.75‖ ISBN 0-8212-6205-X / 978-0-8212-6205-4 Feinstein‘s‘ assemblage of 100 brilliant, captivating, and exquisite photographs make Orchidelarium one of the premier orchid ‗coffee table‘ books available on the market today. The large format and studio photography of primarily single flowers in a variety of views are multidimensional and jump right off the page. At the conclusion the author presents several early photographs take out of doors against a brilliant sky. The effect is quite different but equally as pleasing as the studio shots. Although clearly stated that the book is not intended to be a botanical account of orchids Bob Hesse‘s Introduction is fascinating reading on its own. He covers the popular, scientific, and botanical history of orchids and their many uses as well as the mania that has, and still does, so often surrounds this largest of plant families. Proportionately no more expensive than a smaller format full color book, the price tag is well worth it as either (or both) a gift or to grace a personal orchid library. Ironically the only real error found in the book is the incorrect spelling of Orchidaceae as Orchiadea on page 12. On page 5 the use of the word Foreword is incorrect as it is an Author‘s Preface. By definition a Foreword is written by a recognized expert in the

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subject field and usually praises the content of the book and the author. It is never written by the author. One of the curious aspects of the book is that several photos that have the flowers inverted. These are genera/species that normally have the lip in a non-resupinate, or uppermost, position on the flower, but I will attribute this to the artistic design of the book. The names of the individual flowers are very useful and synonyms are given for those genera that have undergone recent revisions--except for the Florida native Prosthechea cochleata that is listed as Encyclia species. Several photos are simply listed as ‗species‘ that could have easily been identified to hat rank. But this is the botanist speaking and he needs to be reminded that the book is a gallery of exception orchid photographs. Orchidelarium is highly recommended and hopefully it will pique the interest of even the casual reader to search out and locate local orchid and native plant societies and learn more about this premier family of plants!

Vanishing Beauties: Native Costa Rican Orchids. Vol. 1. Acianthera-Kegeliella. F. Pupulin and collaborators (18 collaborators which include most of the recognized specialists in Neotropical orchids are listed on pp 408-409). 2005. ISBN 9977-67- 956-8 (Cloth) xxx+421 pp., numerous color photographs, 25×33 cm. Sistema Editorial y de Difusión Cientifíca de la Investigación, Universidad de Costa Rica, San Jose, Costa Rica. The recent publication of yet another large-format spectacular volume on New World orchids is more than welcome. Given the immense popularity of Costa Rica for ecotours and orchid exploration both the information and the photographs in this book will prove to be an excellent resource. Although not all genera are covered, two which stand out are Beloglottis and Habenaria, those that are treated provide some of the highest quality photographs of orchid species in the current literature. Bonnie Boutell, a personal friend

who lived in Costa Rica for many years had this to say after examining the book. “There are

no words sufficient to describe the experience of living where wild orchids grow. To watch the evolution of the ultimate recyler-the rainforest, convert decay into the exquisite beauty of orchids was life changing. In addition to having them on my own property, I could visit nearby Casa Orchideas Botanical Gardens. The MacAllister family has devoted their lives to creating this magical garden filled with the orchids and plants unique to Costa Rica.”

This is the first volume of a series of three and is to be highly recommended and currently available by inquiry form the Lankester Botanical Garden in Costa Rica http://www.jardinbotanicolankester.org/ing/books.html or from OrchidBooks at http://www.orchidsbooks.com/book.asp?id=846. PMB

http://www.jardinbotanicolankester.org/ing/books.html

http://www.orchidsbooks.com/book.asp?id=846

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Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey Paul Martin Brown with original artwork by Stan Folsom 2007 University Press of Florida. 384 pages 6x9 field guide; 331 full color photographs, 99 line drawings, 86 maps, keys for identification; considerable additional informational material $29.95 Paper (Flexibind): ISBN: 0-8130-9780813030340

Do we really need yet another book on the

orchids of the Northeast? As a native orchid enthusiast, my answer is unequivocally yes. Paul Martin Brown‘s intense devotion to morphological detail sets his orchid books apart from most others; he is almost fanatic in recognizing differences within a species. While some systematic botanists may argue that botanical literature is already overflowing with superfluous plant names, others recognize the value in formally documenting the wide range of morphological variation within plant species. And in much the same tradition as M. L. Fernald, Brown does not hesitate to put pen to paper and describe new taxonomic forms, varieties, and hybrids. In this book, I counted 33 infraspecific orchid taxa from the Northeast originally named by Brown during the past 12 years.

Wild Orchids of the Northeast is the 8th in a series of regional field guides of North

American orchids by Paul Martin Brown and published by University Press of Florida. The book is designed for anyone interested in orchid identification, taxonomy, distribution, and conservation, from beginner to expert. It is a complete revision and expansion of Brown‘s previous work on the subject and now includes all of Pennsylvania and New Jersey, a state never before covered in its entirety.

The book covers 79 species and varieties, 88 forms, and 15 hybrids. It includes an

illustrated key to genera, workable keys to all species, extensive lists of infraspecific taxa and full treatment of synonymy, distribution maps, and species accounts extensively illustrated by high quality photographs and line drawings. The 300+ color photographs are a real strength of the book.

Another strength of the book is the wide range of supplementary material, including

an extensive section on where and how to find orchids at eight regional hot spots: 1) The Northwoods: bog, fen, and boreal forest; 2) Connecticut River Valley; 3) Central New York State; 4) Metropolitan Boston; 5) Cape Cod and Eastern Long Island; 6) The Catskills and Poconos; 7) The Pinelands of New Jersey and Cape May; 8) The Central Alleghenies. I also appreciated the well-illustrated section on ―cryptic species, species pairs, and varietal pairs‖.

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The taxonomy and nomenclature followed by Brown closely follows the Flora of North America treatment of the Orchidaceae published in 2002, with the exception of several species usually included in Platanthera. Brown splits out P. nivea, P. clavellata, and P. integra and includes them in Gymnadeniopsis, a transfer originally proposed by Rydberg in 1901 and favorably noted (but not followed) by FNA author Charles Sheviak. Brown continues to recognize P. pallida as a valid species endemic to eastern Long Island, New York, whereas Sheviak placed it in synonomy under P. cristata, but noted its ―distinctive nature‖. Brown corrected mistakes in previous publications such as the occurrence of P. blephariglottis var. conspicua in New York.

The author‘s stated purpose of this book is ―to assist the user in identifying the wild

orchids of much of the northeastern United States. It is intended to be used primarily in the field and is designed for locating information easily while one foot is in the proverbial bog. The photographs have been taken in the field and are intended to illustrate the species as the user will see them. They are neither studio shots nor great works of art, just good diagnostic photos that portray the plants in their habitats.‖ Brown is being too modest in describing his book and he goes way beyond accomplishing his goals. The Northeast is Brown‘s original stomping grounds and he knows the territory as well as, if not better than anyone. He is intimately connected with these orchids and has spent countless hours pursuing and studying them. Fortunately for us, he enthusiastically shares his vast knowledge and experiences.

I recommend this reasonably priced book for anyone interested in our native

orchids. It has been authored by one of the foremost orchidologists of the region and contains a wealth of information beautifully illustrated on high quality paper. No library will be complete without it. Reviewed by Eric Lamont, Honorary Research Associate, Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458.

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COMPANION CD TO ORCHIDS OF MEXICO LAS ORQUIDEAS DE MÉXICO

Miguel A. Soto Arenas, Eric Hágsater, Rolando Jiménez Machorro, Gerardo A. Salazar, Renato Flores and Ivan González.

Photographs by the authors and 47 additional photographers. A digital photographic catalogue with minimal texts in Spanish and brief comments and instructions in both, Spanish and English languages.

The number of orchids today recorded from Mexico has increased in the last years to surpass 1200 taxa (species and subspecies); in addition, some dozens of natural hybrids. This CD includes a catalogue with some 1500 pictures which illustrate 90% of the Mexican Orchids. It is the complement of the book Orchids of Mexico (Hágsater et.al. 2005) where only 450 species were illustrated. Some pictures are the first published for particular taxa. It includes also a checklist with the orchid names accepted at present and also the several hundred synonyms used in the last 55 years (since the publication of Louis O. Williams, The Orchidaceae of Mexico in 1951). Synonyms are directly linked to current names and these to pictures. Voucher specimens or origin of the plants are given for each picture.

The quality of the photographs, the graphic design, user-friendly management, and the cross-references between accepted names, synonyms and pictures have been very welcome for those who have used it.

Price US $50.00 plus shipment costs. Orders directly to: [email protected]; [email protected]

We are able to accept VISA and MASTERCARD credit cards. Charges will be entered in Mexican pesos and they will appear in your bank statement in your local currency. Please do not forget to include your credit card number, expiration date, cardholder’s name and your card code (3 last digits on the back of your card). Checks in US Dollars, drawn to INSTITUTO CHINOIN, A.C. are also accepted.

Please note an update to the following from what was printed in volume 12: 11, 2006.

Orchids of Manitoba Ames, D., P.B. Acheson, L. Heshka, B. Joyce, J. Neufeld, R. Reeves, E. Reimer, and I. Ward. 2005. Native Orchid Conservation, Inc. Winnipeg, Manitoba.

158 pages, 5.5 x 8.5”, full color

photographs, maps; $17.95 CAD Paper

ISBN 0-9734864-0-6. The first full-color provincial orchid field guide to be published. A group effort that has resulted in a workable and usable field guide with excellent photographs and a distinct slant on conservation. Limited use outside of Manitoba; 3 pages of keys for identification. Available through www.nativeorchid.org



http://www.nativeorchid.org/

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ORCHIDS IN HAWAII Ted Green $10.95 paper 6" x 9" 130 pp. 175+ color photographs ISBN: 1-56647-720-4 Mutual Publishing, LLC

Ted Green

ORCHIDS OF EUROPE, NORTH AFRICA AND THE MIDDLE EAST Pierre Delforge 2006 Timber Press Full color hardcover, 640 pp., 5 x 7.5 in (190 x 125 mm); 1270 color photos plus several line drawings and watercolors ISBN-13:9780881927542; ISBN-10: 0881927546 $39.95



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ORCHIDELIRIUM Harold Feinstein with an Introduction by Robert H. Hesse Bulfinch Press February 2007 $50.00 144 pages 100 full color photographs 13.5 x 11.75‖ ISBN 0-8212-6205-X / 978-0-8212-6205-4

VANISHING BEAUTIES: Native Costa Rican Orchids Vol. 1. Acianthera-Kegeliella. F. Pupulin and collaborators (18 collaborators which include most of the recognized specialists in neo-tropical orchids are listed on pp 408-409). 2005. ISBN 9977-67- 956-8 (Cloth) xxx+421 pp., numerous color photographs, 25×33 cm. Sistema Editorial y de Difusión Cientifíca de la Investigación, Universidadde Costa Rica, San Jose, Costa Rica. Ordering information from: [email protected]

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In Memoriam – T. Sanders McMillan IV (1974-2007)

Jim Fowler

I met the young man in October of 2005 after he had written Paul Martin Brown in hopes of identifying a plant that he believed was an orchid. He had seen Paul‘s book covering Florida‘s orchid species, and hoped that Paul could help him with the identification. Knowing that I live in South Carolina, and that I could possibly visit the site, Paul gave Sanders my email address and encouraged him to contact me.

Sanders was a husband, father of a young son, avid outdoorsman, hunter, and lover of all things in nature. In addition, he had a job that consisted of checking out sites to study the possible environmental impact of proposed development in the immediate vicinity. What Sanders had seen that day in October was several plants, each with four or five lanceolate leaves and a narrow flower spike that had produced several seed capsules. All flower parts were long gone.

Both Paul and I were somewhat stumped, and suggested several possibilities – including (and hoping for) Habenaria quinqueseta. We finally decided that waiting for next year‘s bloom cycle was the only possibility for an accurate identification.

In June of 2006, Sanders sent us several photos that allowed us to accurately pin down the identification to Platanthera lacera – a new species for Richland County, South Carolina. He was quite pleased with his find, and contacted the curator of the herbarium of the University of South Carolina in nearby Columbia, South Carolina. This resulted in the site being set aside as a protected area by the South Carolina Department of Natural Resources – the bureau that is responsible for protecting native flora and fauna in South Carolina. He was right to be proud of his find.

On April 3, 2007, Sanders took his own life. I guess his reason for this exquisitely sad act is not important. Paul never had the opportunity to meet Sanders, and I met him only once, but for some unknown reason, his death has left us both with an unexplained void.

Today, a day after learning of Sanders‘ death, I visited the site once more. It is a small, tree-filled bottomland that most people wouldn‘t give a second look. But today, I found more than twenty of Sanders‘ ―mystery plant‖ in full bloom.

Mind you, I‘m not a firm believer in the after life, but maybe, just maybe, one of these days, I‘ll have another chance to meet this special young man… ―in the stars, my friend."

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ON THE REPORTS OF SPIRANTHES VERNALIS Engelmann & Gray FROM NEW MEXICO

William F. Jennings and Charles J. Sheviak

From time to time, there are literature reports that Spiranthes vernalis is or may be in New Mexico; however, this has been doubted by most authors. Luer (1975) states that "Records of its occurrence in such areas as Ontario and New Mexico are possibly due to confusion with other species." The reports of its occurrence in New Mexico seem to have originated with Ames (1905), who listed Spiranthes vernalis for New Mexico based on a specimen collected by Augustus Fendler, cited by Ames as: "New Mexico, prairies, August 14, 1847." Augustus Fendler was a friend of George Engelmann, and apparently it was through the intervention of Engelmann and Asa Gray that Fendler was allowed to accompany a military detachment from Fort Leavenworth, Kansas to Santa Fe, New Mexico (Ewan, 1950). Fendler traveled via the Santa Fe Trail in 1846, following closely behind General Stephen Watts Kearny, who raised the American flag in Santa Fe on August 18, 1846, completing the bloodless conquest of New Mexico during the Mexican War. Fendler spent the winter in Santa Fe, collected extensively in the spring and summer of 1847, and then returned to St. Louis in the fall of 1847, again via the Santa Fe Trail. His collections were evaluated by Gray (1849) and numerous new species were described. Fendler's collections were some of the earliest specimens taken in the Southwest, and Santa Fe is the type locality for a large number of species. The Santa Fe Trail was used extensively after 1821 to move trade goods by wagon to Santa Fe. By 1843, about 450,000 pounds of merchandise was carried over the trail in 230 wagons (Brown, 1988). When the railroad reached Santa Fe in 1880, the trail became a part of Southwest history. In its heyday, there were actually two trails. Although there were several points of departure in the Kansas City area, since Fendler was traveling with the military, he started at Fort Leavenworth. The trail cut more or less diagonally across Kansas to the Arkansas River at Great Bend. West of Fort Dodge, the trail split. The southern branch (Cimarron Cutoff) continued southwest across the extreme southeast corner of Colorado and the tip of the Oklahoma panhandle. The northern route (Mountain Branch) followed the Arkansas River upstream past Bent's Fort to the mouth of Timpas Creek, then angled southwest to Trinidad, Colorado, and crossed Raton Pass. The trail then followed the base of the mountains, rejoining the Cimarron Cutoff near today's Watrous, New Mexico. The trail then went through Las Vegas, San Miguel, and Pecos, New Mexico, almost exactly on the route of today's I-25. There were no towns along the trail beyond extreme eastern Kansas until Las Vegas, New Mexico was reached, a scant 70 miles from Santa Fe. Fendler traveled westbound via the northern route (Mountain Branch) of the trail in the period August 14 to October 11, 1846. Fendler traveled eastbound via the southern route (Cimarron Cutoff) in the period August 9 to September 24, 1847.

Jennings & Sheviak: ON THE REPORTS OF SPIRANTHES VERNALIS IN NEW MEXICO

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Based on Ames' report, it would appear that Fendler collected the specimen somewhere between Las Vegas and Wagon Mound, New Mexico. However, this is not the case. While the overwhelming majority of Fendler's collections were taken in 1847, some were taken on the trip west in 1846, and this is one of them. The specimen to which Ames referred is at AMES (AMES 71825; GH 1741). It is indeed Spiranthes vernalis, and was annotated so by Ames in 1905, by Correll in 1946, and by Sheviak in 1990. Fendler's original handwritten col-lection label is with the specimen, and clearly states "14/8 46, prairie, no. 32." The specimen was taken on August 14, 1846, one year prior to the date cited by Ames and was taken the first day of the westbound trip, near Fort Leavenworth, Kansas. There is a printed label on the specimen, provided either by Engelmann or Gray, which reads "Plantae Novo-Mexicanae, no. 841 [the number handwritten], A. Fendler coll., 14 Aug [the day and month handwritten], 1847 [printed]. It was common to use a standardized printed label, then to annotate by hand the pertinent data. It was also common in the nineteenth century for herbarium curators who distributed large collections of specimens, such as Fendler's, to renumber the specimens in taxonomic sequence. Thus it was common to see two numbers on old specimens: the collector's number and the curator's number. Additional support for the 1846 collection date comes from Shaw (1982). Shaw reported on Fendler's collection list. Under Orchidaceae, the collection locality of number 841 is given as "low prairies, 9 miles south of Fort Leavenworth; also about 100 degrees west longitude, not far from the bed of the Arkansas River; flowers white.‖ Shaw gives the collection period as between August 14 and September 27, 1846. Ames (1905) reports that Fendler also collected Spiranthes cernua at: "low prairies, 9 miles south of Fort Leavenworth, Fendler 119." Fendler likely numbered his collections as he made them (in chronological order), so Fendler 32 (Spiranthes vernalis) very likely was collected before Fendler 119 (Spiranthes cernua, according to Ames). It appears most likely that Fend-ler collected the Spiranthes cernua along the Arkansas River at the end of August or in early September and not near Fort Leavenworth. Likely, it is the second specimen cited by Shaw. This position also makes sense from a plant geography and blooming season perspective. Very few collections of Spiranthes vernalis are from west of the 98th meridian (roughly a line from Oklahoma City through Wichita and Salina) and the plant is unknown west of the 100th meridian. Based on Kansas specimens cited by Magrath (1971), Spiranthes vernalis blooms in Kansas from June through mid-August, with an average date of July 29 for the specimens cited. Latest collection date cited was August 21. Fendler collecting Spiranthes vernalis in northeastern Kansas on August 14 is a reasonable proposition. Again based on Magrath (1971), Spiranthes cernua (including Spiranthes magnicamporum, since it was not described until 1973) is known from Kansas as far west as 99 degrees 30 minutes. Fendler 119 could have been collected along the Arkansas between Great Bend and Larned. Magrath cited one collection as early as August 27, but all others are after September 13. Average collection date is October 1. Fendler reached Fort Dodge about September 6, so the Spiranthes cernua specimen would have been collected in the first week of September, early for the species, but not impossible. It is also possible that Ames misidentified the specimen, which was not personally seen by me. In summary, reports of Spiranthes vernalis in New Mexico are incorrect, in our opinion, and are based on Ames' erroneous citation of Fendler 32 (841) which was collected in eastern Kansas on


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August 14, 1846, not 1847. A second white-flowered orchid specimen (Fendler 119), considered by Ames to be Spiranthes cernua, was collected in Kansas as well. REFERENCES CITED Ames, O., 1905. Orchidaceae, Fascicle I, A synopsis of the genus Spiranthes north of Mexico, pp.122-156. Brown, W. E., 1988. The Santa Fe Trail. St. Louis: The Patrice Press. Ewan, J.A., 1950. Rocky Mountain Naturalists. Denver: The University of Denver Press. Gray, A., 1849. Memoirs of the American Academy of Arts & Sciences, v. 4, part 1 Luer, C.A. 1975. The Native Orchids of the United States and Canada, excluding Florida. New York: New York

Botanical Garden. Magrath, L.K. 1971. Native Orchids of Kansas. Transactions of the Kansas Academy of Science 74: 287-309. Magrath, L.K., 1973. The Native Orchids of the Prairies and Plains Region of North America, Ph.D. dissertation,

University of Kansas, Lawrence. Shaw, E.A. 1982. Augustus Fendler's collection list, 1846-1847. Contrib. Gray Herb. 212: 1-70. Standley, P.C., 1910, Type Localities of Plants First Described from New Mexico, Contrib. U.S. Nat.

Herb. 13: 143-228. William F. Jennings, P.O. Box 952, Louisville, CO 80027 – [email protected] Charles J. Sheviak, Division of Research and Collections, New York State Museum, Albany, NY 12230 – [email protected]


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NEW FROM THE UNIVERSITY PRESS OF FLORIDA Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey Paul Martin Brown with original artwork by Stan Folsom 2007 University Press of Florida. 384 pages 6x9‖ field guide and more; 331 full color photographs, 99 line drawings, 86 maps, keys for identification; considerable additional informational material $29.95 Flexibind: ISBN: 0-8130-9780813030340

Order from your favorite bookseller, www.upf.com, or signed and inscribed from the authors at [email protected]

http://www.upf.com/


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Don’t Miss Out on Ordering Your Copy of

Orchids of the Dominican Republic

and Haiti

by

ELADIO FERNANDEZ, JUAN LLAMACHO & JAMES ACKERMAN

Available October 2007

220 pages, 100 illustrations

Limit ed pr int ing o f 2000 copies

100 spec ies among the 335 found on the is land.

8.25” x 10.86”

$95 US plus shipp ing at cost

Order ing info r mat ion:

Orders must be p laced by August 15, 2007 at [email protected]

Very few copies are left for this 1st edition so place your order now!

Th e book i s over 200 pa ges , m os t l y col or ph otogr a ph s by Na t ur e Ph ot ogr aph er , El a di o

Fern an dez , a ser i es of es sa ys c o -a u th or ed by J uan Llam a ch o (a uth or of Orc h i ds of Cuba )

an d Ja m es Ackerman (a uth or of Orc h i ds o f Puer t o R i co ) . Th e book a l so in cl udes an

upda t ed l i s t of a l l th e or ch i d speci es r epor t ed for th e i s lan d. Am on g th e or ch i d

ph ot ogra ph s th er e ar e se l dom seen , n ew speci e s an d h a bi t a t ph ot os in bot h coun tr i es . You

m a y vi e w s om e o f th e spect a cu l ar ph ot os a t http://www.gigasize.com/get.php/-

1100139986/070105_Presentacion_Orquideas2.pdf

Spon sor ed by t h e Am er i can Cha m ber of Com m er ce of t h e Dom in i can Republ i c wi t h a

per cen ta ge of th e sa l es devot ed t o AM CHAM' s soci a l r espon si bi l i t y pr oje ct s .


http://www.gigasize.com/get.php/-1100139986/070105_Presentacion_Orquideas2.pdf

http://www.gigasize.com/get.php/-1100139986/070105_Presentacion_Orquideas2.pdf

Documents

June 2007 North American Native Orchid Journal