Native Orchids Research Articles

A. John, An enthusiasm for orchids: sex and deception in plant evolution

Spring arrives in western Australia by the end of August and the shrubby vegetation is colorful and richly perfumed. Some native orchids, though, are brownish or dull-maroon in color, the human nose fails to pick up any fragrance. The strange shapes of the flowers give them such common names as warty hammers (Drakaea), flying ducks (Paracaleana), lazy spiders (Caladenia) and elbow orchids (Arthrochilus). While we smell nothing, these orchids release odors irresistible to many species of wasps. The winged males mistake sculptures and patterns on the lip petals for the bodies of virginal and wingless females. As a lovesick insect backs out of the flower, a plug of sticky pollen globs is deposited on his back. If the same wasp visits a second flower of the same species then cross-pollination occurs and seeds are fertilized although the deceived insect fathers no offspring during this process. More than 40 orchid species found in Western Australia fool male wasps. Why is pollination-by-deceit such a common ‘‘theme’’ in this part of the world? Why not ask Professor John Alcock of Arizona State University? He is known for his research in insect behavior and some past work took him to the southwestern corner of Australia. If you have read his past books such as In a Desert Garden and The Kookburra’s Song you know he can turn the study of evolution into an entertaining and enriching trip. The book is beautifully illustrated with close up photos of the deceptive flowers ‘‘at work.’’ Yes, you will see wasps humiliated by plants within one of the world’s most exciting hotspots of biodiversity. For any serious environmentalist, the chapter on conservation is disturbing. Past agricultural practices in western Australia have reduced available sites where rare species once flourished. They have also introduced European weeds, contributed to a saltier water table and spread an epidemic of cinnamon fungus (Phytophora), killing native species in its path. However, the greatest pleasure in Dr. Alcock’s book is his ability to relate his personal orchid rambles to the work of Charles Darwin. Many of us forget that Darwin turned to orchid pollination following the publication of his Origin of Species. He used orchids as ‘‘model systems,’ in two editions of a book he first wrote in 1862, so the general reader could understand how flower organs fused together and evolved adaptations to better exploit the insects that visit them and transfer their pollen. Consequently, Alcock’s book is most convincing as a study on the survival of the reproductively fit. There’s a nice, early chapter on how and why plants move (also indebted to some of Darwin’s studies on carnivorous and Cassia plants). This is particularly useful for readers, as some Australian orchids have jointed and cantilevered lip petals that flip and reposition male wasps before they have the opportunity to recognize the floral fraud. Let me emphasize, though, that this is not a pretty little adventure in nature appreciation. You will be asked to think early and often and this means reading slowly and thoughtfully. Alcock is one feisty author. He wrestles with the papers of Stephen J. Gould and Richard Lewontin that downplayed the role of adaptation in evolution. For his next trick, Alcock takes a healthy crack at the Creationist movement for still insisting that orchids are beautiful examples of plants created by a sudden act of ‘‘intelligent design.’’ Curiously, Alcock’s book convinced me that both reputable scientists, like Gould and Lewontin, presented arguments as flawed as the stuff published by the people who currently hide under the banner of Creationism. The simple, take-home message from this American in Australia is that, if you ignore the results of almost 160 years of published research and do not bother to P. Bernhardt (&) Department of Biology, St. Louis University, St. Louis, MO 63103, USA e-mail: bernhap2@slu.edu

Native Orchids of the Southern Appalachian Mountains

Native Orchids of the Southern Appalachian Mountains

Thysanoptera Inhabiting Native Terrestrial Orchids in Northern Florida and Southern Georgia

A survey was conducted to identify the thrips species associated with native terrestrial orchids in northern Florida. The above-ground plant structures were sampled when each species was flowering yielding adults of polyphagous species of the common flower thrips Frankliniella tritici (Fitch), Frankliniella fusca (Hinds), and Microcephalothrips abdominalis (Crawford). There is little indication that the orchids were reproductive hosts of these species. Other species of thrips that are more host-specific to orchids were collected and included adults of the orchid-living species Aurantothrips orchidaceus (Bagnall) and Pseudothrips beckhami Beshear and Howell. The presence of larvae of P. beckhami indicated the suitability of at least some orchid species as reproductive hosts. The adults of 2 predatory species in the Family Phlaeothripidae also were collected, Leptothrips mali(Fitch) and Karnyothrips melaleucus (Bagnall).

Wild orchids of the prairies and Great Plains region of North America

Wild orchids of the prairies and Great Plains region of North America

Diversity of mycorrhizal fungi of terrestrial orchids: compatibility webs, brief encounters, lasting relationships and alien invasions

The diversity of mycorrhizal fungi associated with an introduced weed-like South African orchid ( Disa bracteata) and a disturbance-intolerant, widespread, native West Australian orchid ( Pyrorchis nigricans) were compared by molecular identification of the fungi isolated from single pelotons. Molecular identification revealed both orchids were associated with fungi from diverse groups in the Rhizoctonia complex with worldwide distribution. Symbiotic germination assays confirmed the majority of fungi isolated from pelotons were mycorrhizal and a factorial experiment uncovered complex webs of compatibility between six terrestrial orchids and 12 fungi from Australia and South Africa. Two weed-like (disturbance-tolerant rapidly spreading) orchids — D. bracteata and the indigenous Australian Microtis media, had the broadest webs of mycorrhizal fungi. In contrast, other native orchids had relatively small webs of fungi ( Diuris magnifica and Thelymitra crinita), or germinated exclusively with their own fungus ( Caladenia falcata and Pterostylis sanguinea). Orchids, such as D. bracteata and M. media, which form relationships with diverse webs of fungi, had apparent specificity that decreased with time, as some fungi had brief encounters with orchids that supported protocorm formation but not subsequent seedling growth. The interactions between orchid mycorrhizal fungi and their hosts are discussed.

Symbiotic seed germination of Habenaria macroceratitis (Orchidaceae), a rare Florida terrestrial orchid

The rapid loss of native orchid habitat throughout ecologically important areas (e.g., Florida) has prompted researchers to develop appropriate plans for the propagation and reintroduction of many native orchid species. Ideally, symbiotic orchid seed germination methods are utilized in the production of orchid seedlings to be used in plant reintroduction programs. In the current study we (1) describe an efficient symbiotic seed germination protocol to germinate seeds of the rare sub-tropical terrestrial orchid Habenaria macroceratitis; (2) discuss the in vitro fungal specificity demonstrated by this species; and (3) describe the effects of three photoperiods (0/24 h, 16/8 h, 24/0 h L/D) on in vitro symbiotic seed germination of H. macroceratitis. Six fungal mycobionts were isolated from both vegetative and flowering plants of H. macroceratitis from two geographically distinct sites. Symbiotic seed germination percent was highest (65.7%) and protocorm development was most advanced (Stage 2) when seeds were cultured with fungal mycobiont Hmac-310. Seeds of H. macroceratitis demonstrated a degree of specificity toward fungal mycobionts isolated from plants originating from the same site where seed was collected. Continual darkness (0/24 h L/D) inhibited initial seed germination (Stage 1; 17.1%), but stimulated subsequent protocorm development (Stage 2; 53.5%). These findings will aid in developing an efficient symbiotic seed germination protocol for the conservation of this rare Florida terrestrial orchid, and may prove useful in the conservation of other sub-tropical terrestrial orchid species.

Asymbiotic seed germination and in vitro seedling development of Habenaria macroceratitis (Orchidaceae), a rare Florida terrestrial orchid

Continuing loss of native orchid habitat has lead to an increased emphasis on orchid conservation. Major obstacles in the production of native orchid seedlings for use in conservation have been: (1) development of efficient and reliable seed germination protocols and (2) an understanding of early seedling growth and development. Effects of six asymbiotic media (Modified Lucke, Murashige & Skoog, Lindemann, Vacin & Went, Malmgren Modified, Knudson C), four exogenous cytokinins (BA, Zea, Kin, 2-iP), and three photoperiods (0/24, 16/8, 24/0 h L/D) were examined on seed germination and early protocorm development of Habenaria macroceratitis, a rare native Florida terrestrial orchid. Finally, the effects of three photoperiods (8/16, 12/12, 16/8 h L/D) on in vitro seedling development were examined. Percent seed germination was highest on both LM and KC after seven weeks culture (LM = 89.1%, KC = 89.2%); however, protocorm development was enhanced on MM after both seven and 16 weeks. Both zeatin and kinetin at 1 µM enhanced seed germination (Zea = 58.1%, Kin = 47.2%). Final percent seed germination (91.7%) and protocorm development (Stage 4) was increased in the absence of light (0/24 h L/D). In␣vitro seedlings cultured under 8/16 h L/D conditions produced the highest number of tubers per seedling (1.06) with the greatest tuber (42.7 µg) and shoot (fwt = 69.5 µg) biomass and tuber diameter (3.1 mm).

In vitro seed culture and seedling development of Calopogon tuberosus

A major obstacle to native orchid production is difficulty in seed germination. Culture media and light effects on seed germination of Calopogon tuberosus var. tuberosus, a native orchid with horticultural potential, were studied. Culture media included Knudson C, Malmgren modified terrestrial orchid, and PhytoTechnology orchid seed sowing. Effects of 8 weeks continual darkness, 8 weeks 16-h photoperiod, 2 weeks dark followed by 6 weeks 16-h photoperiod, 4 weeks dark followed by 4 weeks 16-h photoperiod, and 6 weeks dark followed by 2 weeks 16-h photoperiod were examined. Percent seed germination was highest on Knudson C after 8 weeks culture; however, seedling development was enhanced on PhytoTechnology seed sowing medium during 8 weeks culture under a 16-h photoperiod. This suggests that while KC and darkness promoted seed germination, P723 and light enhanced further seedling development. Seedlings of C. tuberosus readily acclimated to greenhouse conditions.

Survival of transplanted terrestrial orchid seedlings in urban bushland habitats with high or low weed cover

The conservation of wild orchid populations may depend on the establishment of propagated orchids to field sites to help sustain depleted populations if natural recruitment is not successful. However, very little is known about biotic factors which influence the establishment of terrestrial orchid seedlings in natural habitats. An experiment was established to measure the survival of six orchid species during their first growing season following transplantation to a West Australian urban bushland with a Banksia and Eucalyptus canopy and understorey dominated either by weeds or native vegetation. Symbiotically germinated orchid seedlings raised in the laboratory for 5 months before planting were established in adjacent field sites with high or low weed cover. There was a gradual mortality of seedlings at field sites throughout the growing season, primarily owing to insect grazing, and this was not affected by the enclosure of seedlings by wire mesh or shade cloth. Overall rates of survival varied from 49% for Microtis media R.Br., a species capable of growing in disturbed habitats, to 21% for Caladenia arenicola Hopper & A.P.Brown, the most common native orchid at these sites. However, not all surviving seedlings produced a tuber, so their expected rate of survival after the next dry season was reduced further. The factors having the greatest impact on seedling survival were site aspect (slope and canopy cover), weed cover and orchid species respectively. Orchid seedling survival was not well correlated with the presence of existing orchids of the same species at the same sites or the presence of compatible fungi in soil at these sites (simultaneously measured by orchid seed baiting).

Fine scale distribution of pollinator explains the occurrence of the natural orchid hybrid × Orchis bergonii

:Hybridization is frequent in Orchidaceae and may play an important role in their speciation. However, information about pollinators and their role in hybridization is sparse. In the Mediterranean region of France, confirmed pollinatorsobserved on Orchis simia and O. anthropophora included only one species common to both, the beetle Cidnopus pilosus. This beetle thus appears responsible for hybridization. Cross-specific pollen transfer is bidirectional, but hand pollination showed higher fruit set after deposition of O. anthropophora pollen on the stigma of O. simia than in the opposite direction. Both locally and over a 1600-km2 study area, the presence of hybrids was shown to be strongly predicted by the microhabitat preferences of this beetle. To my knowledge, this is a novel demonstration that the spatial distribution of a natural hybrid in plants can be predicted by the local occurrence of a pollinator species identified as effecting hybridization. I suggest that similar interaction could explain the distribution of hybrids in other orchid species pairs.

Substrates and fertilization for the rustic cultivation of in vitro propagated native orchids in Soconusco, Chiapas

AbstractNative orchid cultivation is a compatible alternative for impoverished coffee farmers in southeastern Mexico who are in crisis due to falling prices. Sustainable orchid cultivation is also an urgent necessity as an alternative to nonsustainable extraction from protected reserves, forest fragments and traditional coffee plantations, and to restore and conserve populations within these habitats. Our objective was to test the effects of locally available substrates and fertilizers upon orchids cultivated under typical rural conditions in coffee-producing areas in Soconusco, Chiapas. Seven species of epiphytic orchids native to Soconusco region—Cattleya aurantiaca, Brassavola nodosa, Prosthechea (Encyclia) chacaoensis, Anathallis (Pleurothallis) racemiflora, Cattleya skinneri, Cycnoches ventricosumandEncyclia cordigera—were propagatedin vitro, acclimatized and established in rustic orchid galleries in the home gardens and plantations of coffee growers. Locally available waste products were used as substrates: clay tiles, tree bark, bamboo, seed hulls of pataxte (Theobroma bicolor) and wire baskets filled with bark chips. Two cheap and readily available commercial foliar feeds, Algaenzims (an organic product) and Bayfolan (a synthetic product) were tested. First, the substrates alone were tested for a period of 6 months to 1 year, then a combination of substrates and fertilizers were tested for 6 months, for effects upon leaf and root growth and root number. The mortality rates of these nonsymbiotically propagated, epiphytic orchids during the acclimatization phase, prior to these experiments, were high, between 60 and 90%. Once established in rustic galleries, the young orchid plants showed no preference for a particular substrate, survival depended upon technical problems during establishment, relating to difficulties with the attachment of plants to substrates, and the variable quality of care and attention offered by the farmers. Both fertilizers significantly improved one or all the parameters studied, and possibly counteracted the negative effects of the absence of symbiotic fungi, which, under natural conditions, are essential for orchid seed germination and adequate development of the young plant. More than half of the producers did not continue with the orchid cultures for economic and cultural reasons.

ACTIVATED CHARCOAL FOR IN VITRO PROPAGATION OF BRAZILIAN ORCHIDS

This research focused on the effects of activated charcoal on the in vitro growing media used in the propagation of Brazilian native orchids. Activated charcoal has been described improving the in vitro plant growing and survival after transplant. Seeds were germinated in vitro and after three months they were transferred to one of the following media: MS (Murashige and Skoog) media; half strength of MS media; MS media supplemented with 1 g.L of activated charcoal; half strength of MS media supplemented with 1 g.L of activated charcoal; MS media supplemented with 2 g.L of activated charcoal; half strength of MS media supplemented with 2 g.L of activated charcoal. After six months the following features were evaluated, plantlet height, number of branches per plant, number of roots per plant, length of the roots, fresh weight and O. trulliferum was also evaluated concerning with the pseudobulb diameter. Just after the in vitro plant features evaluation they were transferred to pots containing the tree fern fiber and sphagnum. Two months after transplant the survival rate was evaluated for each treatment. Activated charcoal improved the in vitro plant quality and increased the survival after transplant of the three species evaluated. To M. flavescens and L. flava best results were obtained with half strength of MS media supplemented with 2 g.L of activated charcoal and O. trulliferum with MS media supplemented with 1 g.L of activated charcoal. INTRODUCTION Orchids are one of the most valuable and appreciated plants in the world. Due to its commercial value and beauty, orchids are very cultivated. Several species are in risk of extinction due to its predatory collection and destruction of its habitat. To preserve orchids using tissue culture technique is an option to produce orchids in large scale (Silva, 1986). An adequate formulation of the nutrition media is essential to tissue culture, because media must supply the essential substances for in vitro growth development (Caldas et al., 1990). Addition of activated charcoal on tissue culture media can be beneficial or adverse to growth and development. Its presence depends on the media, the specie and the tissue used and/or the objectives of the research (Pan and Staden, 1998). Activated charcoal as supplement on the media has been reported through the years by several authors due to its beneficial effects, such as, absorption of phenolic complex (Pan and Staden, 1998); root stimulator (George and Havishankar, 1997; Caldas et al., 1990); rhizome growth (Kim and Lee, 1992); development improvement (Choi and Chung, 1989); and absorption of toxic substances present in the media (Fridborg and Eriksson, 1978). Waes (1987) used activated charcoal for in vitro development of 18 species of European native orchids with concentrations from 0,02 % to 0,03 % (g.L) to improve in vitro development. The MS media contains nitrogen in ammonium and nitrate forms. Vitrification is frequent observed affecting plantlets development. Reduction of nitrogen concentration on ammonium has been used to reduce vitrification (Caldas et al., 1990). Reduction of MS media to half strength for in vitro orchid has been showed positive to reduce Proc. V IS on New Flor. Crops Eds.: A.F.C. Tombolato and G.M. Dias-Tagliacozzo Acta Hort. 683, ISHS 2005

Propagação <em>in vitro</em> de orquideas brasileiras utilizando meios de cultura tradicionais e formulações com fertilizantes comerciais

Some species of Brazilian orchids are practically extinct in their natural habitats. Large scale propagation of in vitro native orchids is a way of preserving them. Specific culture media for some species have already been described, but there are no studies on Catasetum fimbriatum and Cyrtopodium paranaensis. The aim of the present study was to establish in vitro propagation protocol of C. fimbriatum and C. paranaensis to reintroduce these species in natural habitat. The traditional culture media were: MS; MS modified with ½ macronutrients; MS modified with ¼ macronutrients, Vacin & Went and Knudson C. The commercial fertilizers formulations were: N.P.K (10-5-5) 2 mL L-1 and formulation N.P.K (10-30-20) 3 g L-1. The data, assessed six months after the beginning of the experiment, consisted of: number of roots, main root length, pseudobulb diameter, plant dry weight, root dry weight and plant height. The best results of C. fimbriatum plant development were obtained using MS medium complete and MS medium modified with half of macronutrients for canopy length. For the number of roots, the best results were obtained using MS modified with half of macronutrients. The best results for C. paranaensis plant development were obtained using MS modified with half of the macronutrients for canopy length and MS modified with ¼ of macronutrients for number of roots.

Isolamento e identificação de fungos micorrízicos rizoctonióides associados a três espécies de orquídeas epífitas neotropicais no Brasil

Distúrbios causados pelo homem têm resultado no aumento do risco de extinção de diversos táxons de orquídeas nativas da Mata Atlântica no Brasil. Na natureza, orquídeas utilizam obrigatoriamente fungos endomicorrízicos para a germinação de sementes e desenvolvimento da plântula, ao menos nos primeiros estádios do seu ciclo de vida. Assim, fungos micorrízicos associados ao sistema radicular de orquídeas nativas vêm sendo isolados, caracterizados e armazenados para uso em futuros programas de conservação de espécies de orquídeas, por meio da germinação simbiótica. Três isolados de fungos micorrízicos rizoctonióides foram obtidos do sistema radicular de três espécies de orquídeas neotropicais, Gomesa crispa, Campylocentrum organense e Bulbophyllum sp., de três diferentes fragmentos de Mata Atlântica no Brasil. Estudos taxonômicos, baseados na condição nuclear, morfologia da hifa vegetativa e ultra-estrutura do septo dolipórico, revelaram que os isolados pertencem aos gêneros Ceratorhiza e Rhizoctonia. Esse é o primeiro relato do isolamento de fungos micorrízicos associados ao sistema radicular dessas espécies de orquídeas neotropicais. Aspectos relativos à taxonomia e ao uso desses isolados no contexto de um programa de conservação de orquídeas nativas são discutidos.

Distribution of crassulacean acid metabolism in orchids of Panama: evidence of selection for weak and strong modes.

Crassulacean acid metabolism (CAM) is one of three metabolic pathways found in vascular plants for the assimilation of carbon dioxide. In this study, we investigate the occurrence of CAM photosynthesis in 200 native orchid species from Panama and 14 non-native species by carbon isotopic composition (δ13C) and compare these values with nocturnal acid accumulation measured by titration in 173 species. Foliar δ13C showed a bimodal distribution with the majority of species exhibiting values of approximately -28‰ (typically associated with the C3 pathway), or -15‰ (strong CAM). Although thick leaves were related to δ13C values in the CAM range, some thin-leaved orchids were capable of CAM photosynthesis, as demonstrated by acid titration. We also found species with C3 isotopic values and significant acid accumulation at night. Of 128 species with δ13C more negative than -22‰, 42 species showed nocturnal acid accumulation per unit fresh mass characteristic of weakly expressed CAM. These data suggest that among CAM orchids, there may be preferential selection for species to exhibit strong CAM or weak CAM, rather than intermediate metabolism.

Orchid species succession in rehabilitated bauxite mines in Western Australia

Twenty-three orchid species were recorded in Alcoa's permanent vegetation-monitoring plots in unmined and rehabilitated jarrah forest. Of these, 22 were identified in the unmined jarrah forest and 20 were recorded in rehabilitated areas of between 1 and 31 years old. Three species (Cyrtostylis ovata, Lyperanthus serratus and Prasophyllum elatum) were only recorded in the unmined forest and one species was only recorded in rehabilitated areas (Diuris carinata). The overall density of native orchids in the forest was 13 755 plants ha–1, 10 times greater than the density in rehabilitated areas (1381 plants ha–1). The most abundant species in the forest were Cyrtostylis robusta, Caladenia flava, Pterostylis nana and Thelymitra crinita, all with densities greater than 1000 plants ha–1. The most abundant species in the rehabilitated areas were Microtis media, Disa bracteata (an introduced species), Caladenia flava, Pterostylis nana, Diuris longifolia and Pterostylis vittata, all with densities greater than 60 plants ha–1. In rehabilitation older than 10 years, the density of orchids increased to 2685 plants ha–1. Burning in rehabilitated areas resulted in large increases in orchid densities. It is believed that orchid colonisation of rehabilitated bauxite mines is dependent on symbiotic mycorrhiza, which are in turn dependent on development of an organic litter component in the soil.

Orchid viruses of natural Ukrainian flora

Virus infection greatly affects to the normal growth and reproductive intensity of orchid species in greenhouses. Peculiarities of ornamental orchid viruses has become known in different countries of the world while there is nothing known about orchid virus infection in natural flora of Europe, for example in Ukraine. After extensive virus detection of natural orchids of Ukraine (Carpathians, Crimea), some viruses infecting these plants in their natural inhabitance were determined.

Physiological Characteristics of Symbiotic Fungi Associated with the Seed Germination of Gastrodia elata

This study was carried out to investigate the possibility for seeds germination of Gastrodia elata using symbiotic fungi. Since seeds of G. elata are very small and lack an endosperm and other nutrients, their germination is difficult without requirement for external nutrients. Out of twenty six isolates collected from protocorms of G. elata and roots of native orchids inhabited in wild, two strains (H-2 and H-21) were observed to stimulate the seed germination of G. elata. The seed germination of G. elata was excellent on oak tree leaves medium. The optimal conditions for mycelial growth of symbiotic fungi were 25°C and pH 6.0, respectively. The mycelial growth of H-2 strain was excellent on YMA medium, while H-21 was poor on PDA medium. In case of carbon sources, the mycelial growth of H-2 and H-21 was good on media supplemented with glucose and dextrin, respectively. Calcium nitrate was good for mycelial growth of H-2 strain as a nitrogen sources, whereas urea was effective to H-21 strain.

(1481) Proposal to conserve the name Dactylorhiza Necker ex Nevski over Coeloglossum Hartm. (Orchidaceae)

The orchid genera Dactylorhiza and Coeloglossum are among the most widely distributed of all north temperate orchid genera, both being reported from Europe, Asia as far south as the Himalayas and in northern North America, e.g., Moore & So6 (in Tutin & al., Flora Europaea 5: 335-337. 1980), Stace (New Flora of the British Isles ed. 2: 973-979. 1997), Renz & Taubenheim (in Davis, Flora of Turkey 8: 450-599. 1984), Renz (in Rechinger, Flora Iranica 126: 58-139. 1978), and Luer (Native Orchids of the United States and Canada: 154-160, 170-174. 1975). Dactylorhiza species are numerous and prominent and distinctive elements of the vegetation, especially of lowland and montane marshes and meadows; the solitary species of Coeloglossum is rather inconspicuous and found in grassland and on the edges of woods in both lowland and montane habitats. Using molecular and morphological data, recent cladistic analyses of European members of the tribe Orchidoideae (Orchidaceae) by Pridgeon & al. (in Lindleyana 12: 89-109. 1997), Bateman & al. (in Lindleyana 12: 113-141. 1997), and Pridgeon & al. (Genera Orchidacearum 2: 2001) have demonstrated that Coeloglossum nests within the much larger genus Dactylorhiza. In habit, Coeloglossum viride (L.) Hartm. falls within the range of variation of Dactylorhiza but its flowers are green or brown and lack a spur. It interbreeds freely with Dactylorhiza species in the wild and numerous natural hybrids between them have been described. Bateman & al. (l.c.) and Pridgeon & al. (l.c., 2001) consider that they are congeneric and use the name Dactylorhiza for the combined genus. Coeloglossum, however, is the earlier generic name, having been published by Hartmann (Handb. Skand. Fl. ed. 1: 329. 1820) based upon Satyrium viride of Linnaeus (Sp. PI.: 944. 1753) while Dactylorhiza, although coined by Necker in Elementa Botanica iii: 129 (1790), a suppressed work for publication of generic names (Art. 32.7 and App. V), was first validated by Nevski (in Acta Inst. Bot. Acad. Sci URSS. ser. 1, fasc. 4. 1937). Dactylorhiza was lectotypified by Orchis umbrosus Kar. & Kir. (D. umbrosa (Kar. & Kir.) Nevski) by So6 (Orchidee (Hamburg) 19: Sonderheft 19. 1968). The rule of priority in the International Code

Native Orchids of Oklahoma

As of the publication of this paper Oklahoma is known to have orchids of 33 species in 18 genera, which compares to 20 species and 11 genera reported by Waterfall (1969). Four of the 33 species are possibly extinct in the state based on current survey work. The greatest concentration of orchid species is in the southeastern corner of the state (Atoka, Bryan, Choctaw, LeFlore, McCurtain and Pushmataha Counties).