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INTERNATIONAL JOURNAL ON ORCHIDOLOGYISSN 1409-3871VOL. 11, No. 3 DECEMBER 2011PROCEEDINGS OF THE THIRD SCIENTIFIC CONFERENCE ON ANDEAN ORCHIDSEdited by ALEC M. PRIDGEON Royal Botanic Gardens, Kew Richmond, Surrey TW9 3AB, United Kingdom and HUGO GUILLERMO NA V ARRETE ZAMBRANO Apdo. 17-01-2184, Quito, Ecuador
The Vice-Presidency of Research UNIVERSITY OF COSTA RICA is sincerely acknowledged for his support to the printing of this volume
INTERNATIONAL JOURNAL ON ORCHIDOLOGY Copyright 2011 Lankester Botanical Garden, University of Costa Rica Effective publication date: November 20, 2011 Layout: Jardn Botnico Lankester. Cover: Sobralia mucronata Ames & C. Schweinf. Photograph by F. Pupulin. Printer: S&F Printing Solutions Printed copies: 500 Printed in Costa Rica / Impreso en Costa RicaR Lankesterian a / International Journal on Orchidology No. 1 (2001)-. -San Jos, Costa Rica: Editorial ISSN-1409-3871 1. Botnica Publicaciones peridicas, 2. Publicaciones peridicas costarricenses
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T ABLE OF CONTENTS Preface ALEC A. PRIDGEON Acknowledgments Darwins orchids: their variation, plasticity, and natural selection JAMES D. ACKERMAN, MARIELY MORALES & RA YMOND TREMBLA Y How many orchid species in Costa Rica? A review of the latest discoveries DIEGO BOGARN Mating systems in the Pleurothallidinae (Orchidaceae): evolutionary and systematic implications EDUARDO LEITE BORBA, ARIANE RAQUEL BARBOSA, MARCOS CABRAL DE MELO, SAMUEL LOUREIRO GONTIJO & HENRIQUE ORNELLAS DE OLIVEIRA A look at The orchid book in celebration of Charles Darwins 200th birthday KENNETH M. CAMERON Orchids in a changing climate PHILLIP CRIBB Sobralia warszewiczii ROBERT L. DRESSLER Filogenia molecular preliminar de Scaphosepalum (Orchidaceae: Pleurothallidinae) LORENA ENDARA A., NORRIS H. WILLIAMS & W. MARK WHITTEN The genus Coryanthes: a paradygm in ecology GNTER GERLACH What will be left of the primary forest in Ecuador? ALEXANDER HIRTZ How uniform is species diversity in tropical forests? PA VEL KINDLMANN & CARLOS A. VERGARA CASSAS Active mountain building and the distribution of core Maxillariinae species in tropical Mexico and Central America STEPHEN H. KIRBY The role of common orchids in appreciating the complexity of biodiversity conservation MARILYN H. S. LIGHT & MICHAEL MACCONAILL Botanic gardens, education, and orchid conservation strategies: the need for a coordinated approach THOMAS J. MIRENDA Preliminary molecular phylogenetics of Sobralia and relatives (Orchidaceae: Sobralieae) KURT M. NEUBIG, W. MARK WHITTEN, MARIO A. BLANCO, LORENA ENDARA, NORRIS H. WILLIAMS& SAMANTHA KOEHLER Epigenetic information Unexplored source of natural variation OVIDIU PAUN & MARK W. CHASE III IV 179 185 207 223 233 239 245 253 265 269 275 293 301 307 319 LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. I
Of greenish Encyclia: natural variation, taxonomy, cleistogamy, and a comment on DNA barcoding FRANCO PUPULIN & DIEGO BOGARN The leaves got right again: DNA phylogenetics supports a sister-group relationship between Eurystyles and Lankesterella (Orchidaceae: Spiranthinae) GERARDO A. SALAZAR & ROBERT L. DRESSLER Orchid seed stores for sustainable use: a model for future seed-banking activities PHILIP T SEA TON & HUGH W. PRITCHARD Fitness landscapes in orchids: parametric and non-parametric approaches RA YMOND L. TREMBLA Y POSTERS Conservation science Aplicacin de la tcnica de encapsulacin deshidratacin para la crioconservacin de semillas y protocormos de Oncidium stenotis (Orchidaceae) ALBERTO ROURA, KARINA PROAO & MNICA JADN Ecology Mating system and female reproductive success of the endemic, epiphytic Prosthechea aff. karwinskii (Orchidaceae) ERIKA CAMACHO-DOMNGUEZ & IRENE VILADAZ Laelia speciosa ROBERTO GARIBA Y-ORIJEL, KEN OY AMA & IRENE VILA-DAZ Evidence of protandry in Aa Rchb.f. (Orchidaceae, Cranichideae) DELSY TRUJILLO, THASSILO FRANKE & REINHARD AGERER Systematics A phylogenetic analysis of the genus Pleurothallis, with emphasis on Pleurothallis subsection Macrophyllae-Fasciculatae M. WILSON, C. BELLE, A. DANG, P. HANNAN, C. KENYON, H. LOW, T ST A YTON & M. WOOLLEY Phylogenetic analysis of the Andean genus Brachycladium Luer (syn. Oreophilus Higgins MARK WILSON & LOU JOST Barcoding the species of Pleurothallis subsection Macrophyllae-Fasciculatae MARK WILSON LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. II LANKESTERIANA 325 337 349 355365 366 367 368 369 370 371
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. III PREFACE the Orchid Society of Quito, Botanical Garden of Quito, and Municipality of Quito to host the Third Conference It is appropriate that we held the Conference in Ecuador in 2009, the bicentenary of the birth of Charles became aware of morphological differences among them, differences that translated into recognition of differ ent species on each of the islands in the archipelago and would later be adduced to support his theory of natural uador, while Thomas Mirenda and Philip Seaton discuss in situ and ex situ solutions, whether already in progress or proposed. As always, education is the underlying key to all the solutions, and it is in that context we hope this ALEC M. PRIDGEON
ACKNOWLEDGMENTS many sponsors: Sociedad de Orqudeas de Quito, Jardn Botnico de Quito, Municipio de Quito, Ministerio del Medio Ambiente, Ministerio de Turismo, Fundacion Botnica de los Andes, and Poning tours and assistance in producing the incredible Show. Publication of the Proceedings of this ALEC M. PRIDGEONHUGO GUILLERMO NA V ARRETE ZAMBRANO LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. IV LANKESTERIANA
his treatise, On the Origin of Species by Means of Natural Selection. Unquestionably, this was a western society. This 450-page abstract established the process by which change may occur through natural LANKESTERIANA 11(3): 179. 2011.DARWINS ORCHIDS: THEIR VARIATION, PLASTICITY, AND NATURAL SELECTION JAMES D. ACKERMAN 1,4 MARIELY MORALES 2 & RA YMOND TREMBLA Y 31 23 4 Author for correpondence: firstname.lastname@example.org; ABSTRACT cloud the effects of selection? Using a Tolumnia variegata population as a model system, we conducted a likely a consequence of dramatically different rainfall. Fruit production was similar for sun and shade sites, but respond in the same way. RESUMEN Las consecuencias de lo plasticidad fenotpica? Usando poblaciones de Tolumnia variegata en la precipitacin. La produccin de frutos fue similar entre sitios de sombra y sol, pero las trayectorias de seleccin de las caractersticas entre los dos lugares fueron en su mayora incongruentes. Nuestros datos indican caracterst icas responden de la misma manera. KEY WORDS :
was slow to amass supporting data and was pushed to come forth with his thesis when Alfred Russel Wallace wrote to him expressing essentially the same ideas From those studies and many others throughout the unmatched by all those who preceded him and those who followed. enrich it with accumulated knowledge of biology, especially with a better understanding of genetics. cornerstone to the theory thoroughly appreciated by remarkable considering modern genetics had not yet selection ensues with a predictable outcome (Endler, 1986). adaptations that ensured or enhanced the probability for cross-pollination. The amount of data he accumulated the Origin of Species his landmark treatise was On the Various Contrivances by which British and Foreign Orchids Are Fertilised by Insects, a compendium of pollination mechanisms natural selection. to catalogue them, that he also had to show what the of his theory of natural selection: The Effects of Cross and Self Fertilisation in the Vegetable Kingdom exaggeration to say that Nature tells us, in the most emphatic manner, that she abhors perpetual selfOrchids are not only an ideal model system to elucidate the remarkable adaptations associated with cross-pollination, but they are also ideal because they et al of the basic conditions for natural selection. Flowers (Ackerman, 1986). Why do many species of orchids show high isolation mechanisms among closely related species. pollinator-sharing, creating hybrid swarms through habitats that bring two otherwise separated species together. Some well-documented hybrid swarms et alet al., 2006).LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.180 LANKESTERIANA
A second possibility is that periodic genetic orchid populations do appear to be small, and fruit set consistent with this mechanism of generating unusually Pellegrino et al et al., 2007). Genetic drift may also occur in larger populations when the time. Although this has been demonstrated for some Marchand & Ackerman, 2006), we are not aware of a similar example in orchids. (Endler, 1986), but there are types of selection that can a reward to their pollinators who search for food, to attract mates. These come in the form of nectar, on the pollination system (Simpson & Neff, 1983). Thousands of orchid species, though, do not offer any through deception by appearing to offer the resources that pollinators seek. Perhaps the most common form of deceit is the appearance as a food resource (Ackerman, 1986). Precise mimicry is not necessary as certain type, and if they fail to extract a reward, then pollination (Johnson & Nilsson, 1999; Johnson et al., would be better represented in the next generation. plants has been frequently commented upon, but there is only a single published report that compares this was a single species pair of Anacamptis et al. that indicate deception-pollinated plants are indeed one case has been reported -Dactylorhiza sambucina et al et al., 2006). Thus far, all other that relaxed selection or some other form of selection may be occurring (Ackerman et al., 1997; Aragn & Ackerman, 2004; Ackerman & Carromero, 2005). has not or will not occur but that only during the course of these particular studies it was not happening. et al., 2005). Of course, there is yet another explanation: the methods for detecting selection were not rigorous when it occurs. Variation and plasticity Variation is often presumed to be heritable, but the genetics of these traits are rarely examined in orchids, primarily because traditional methods require common For this, orchids are not good model systems because LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. ACKERMAN et al. 181
expression occurs and is not likely directly inherited. and Johnson, 2005). We get around this problem by gradients occur within populations, and plants respond from year to year as habitats go through successional processes or as local climate changes, whether plants (Pigliucci, 2001), yet plasticity in orchids has selection within a population may become additionally problematic. Certainly phenotypic plasticity can blunt the power of selection. Plasticity in orchids: a test case question in a recent paper (Morales et al., 2010) by studying a population of Tolumnia variegata (Sw.) Braem, a deception-pollinated twig epiphyte occurring in the Greater Antilles from the Virgin Islands in the east to western Cuba. This is a species for which dependent selection but without success (Ackerman et al., 1997). In fact, no selection of any kind on any Large populations of T. variegata are not uncommon and can be found from dry forests to wet montane regions (Ackerman, in press). Plants are small with sympodial growth. Morphological and genetic and Ward, 1999). Flowers are self-incompatible, and pollinators are female Centris decolorata (Apidae; Ackerman et al. To induce plastic responses, we conducted a reciprocal transplant experiment where our grew in the open under full sun and also in the shade. We used 80 plants from two sun sites and 80 plants from two shady sites, all of which were within a 25 m radius. Prior to transplanting the orchids, we transplanting, we followed these plants for two years second year (2000) was a dry one, with precipitation approximately 500 mm below normal. results of the transplant experiment, we found that plant responses were complex (Morales et al., 2010). Leaf characteristics of plants transplanted from sun to and stayed in the shade. Likewise, those that went from general was strongly affected by drought conditions in the second year. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.182 LANKESTERIANA FIGURE Tolumnia variegata
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. ACKERMAN et al. 183 was strongly affected by year, whereas the number of Floral characteristics we measured were associated appeared to be plastic, and all were affected by year. plant traits between the sun and shade sites. We did action may occur, as has been shown in other orchid systems (Ackerman & Carromero, 2005). Because the picture thus far is clear only in the approach to detecting patterns of selection by using the non-parametric, cubic spline regression analysis (see Tremblay et al., 2010). We used data for only those repeated measures ANOVAs. We found that for petal for shade and sun plants. Thus, the two habitats mostly limited to sun plants because shade plants is what was detected in an earlier study at the same locality (Sabat and Ackerman, 1996). Finally, we When selection across microhabitats is consistent, are on different trajectories, then the speed of change Conclusions populations may be explained by the possibility that selection is not occurring, or as we discussed here, selection may go in different directions depending on where they are in a heterogeneous habitat. The to year. Using a Bayesian approach with data from this Tolumnia study plus data from a population of Caladenia, Tremblay et al. (2010) found that indeed to detect in orchids. The notion that selection need not be operational all the time should come as no surprise. whereby periods of drift (no selection) may be punctuated by brief but strong bouts of selection (Tremblay et al., 2005). LITERATURE CITED Ackerman, J. D. 1986. Mechanisms and evolution of fooddeceptive pollination systems in orchids. Lindleyana 1: 108-113. Ackerman, J. D. 1989. Limitations to sexual reproduction in Encyclia krugii (Orchidaceae). Syst. Bot. 14: 101-109. Ackerman, J. D. In press. Flora of the Greater Antilles: Orchidaceae. Memoirs of the New York Botanical Garden, Bronx, USA. Ackerman, J. D. & W. Carromero. 2005. Is reproductive success related to color polymorphism in a deceptionpollinated, tropical terrestrial orchid? Caribbean J. Sc. 41: 234-242. Ackerman, J. D. & M. Galarza-Prez. 1991. Patterns and maintenance of extraordinary variation in the Caribbean orchid, Tolumnia (Oncidium) variegata. Syst. Bot. 16: 182-194. Ackerman, J. D. & S. Ward. 1999. Genetic variation in a widespread, epiphytic orchid: where is the evolutionary potential? Syst. Bot. 24: 282-291. variation matter in deception pollinated Psychilis monensis (Orchidaceae)? Oecologia 138: 405-413. Azevedo, C., E. L. Borba & C. van den Berg. 2007. Evidence of natural hybridization and introgression in Bulbophyllum involutum Borba, Semir & F. Barros and
B. weddellii (Lindl.) Rchb. f. (Orchidaceae) in the Chapda Diamantina, Brazil, by using allozyme markers. Rev. Brasil. Bot. 29: 415-421. Darwin, C. 1845. Journal of researches into the natural history and geology of the countries visited during the voyage of H.M. S. Beagle round the world under the command of Capt. Fitz Roy, R. N. John Murray, London, UK. Darwin, C. 1859. On the origin of species by means of natural selection John Murray, London, UK. Darwin, C. 1862. On the various contrivances by which British and foreign orchids are fertilised by insects. John Murray, London, UK. Darwin, C. 1877. The effects of cross and self fertilisation in the vegetable kingdom John Murray, London, UK. Endler, J. A. 1986. Natural selection in the wild. Princeton University Press, USA. Gigord, L. D. C., M. Macnair & A. Smithson. 2001. Negative frequency-dependent selection maintains a orchid Dactylorhiza sambucina (L.) So. Proc. Nat. Ac. Sc. USA 98: 6253-6255. Grant, B. R & P. Grant. 2006. Evolution in Darwins Galpagos Archipelago. Zoology 106: 255-259. variety and blooming times. Evolution 29: 325-334. Heinrich, B. 1979. Bumblebee economics. Harvard University Press, Cambridge, Massachusetts, USA. Herrera, C. M. 2009. Multiplicity in unity. Plant subindividual variation and interaction with animals. University of Chicago Press, Chicago, Illinois, USA. Oecologia 147: 60-68. Jerskov, J., P. Kindlmann & S. S. Renner. 2006. Is the colour dimorphism in Dactylorhiza sambucina maintained by differential seed viability instead of frequency-dependent selection? Folia Geobot. 41: 61-76. Johnson, S. D. & L. A. Nilsson. 1999. Pollen carryover, geitonogamy, and the evolution of deceptive pollination systems in orchids. Ecology 80: 2607-2619. Johnson, S. D., C. L. Peter & J. gren. 2004. The effects of nectar addition on pollen removal and geitonogamy in the non-rewarding orchid Anacamptis morio. Proc. Royal Soc., London B 271: 803-809. Morales, M., J. D. Ackerman & R. L. Tremblay. 2010. gradient in the epiphytic orchid, Tolumnia variegata: 163: 431-446. Pellegrino, G., A. M. Palermo, M. E. Noce, F. Bellusci, & A. Musacchio. 2007. Genetic population structure in the Mediterranean Serapias vomeracea, a nonrewarding orchid group. Interplay of pollination strategy and stochastic forces? Pl. Syst. Evol. 263: 145-157. Pigliucci, M. 2001. Phenotypic plasticity: beyond nature and nurture. The Johns Hopkins University Press, Baltimore, Maryland, USA. Ramrez, S. R., B. Gravendeel, R. B. Singer, C. R. Marshall & N. E. Pierce. 2007. Dating the origin of the Orchidaceae from a fossil orchid with its pollinator. Nature 448: 10421045. Rivera-Marchand, B. & J. D. Ackerman. 2006. Bat pollination breakdown in a Caribbean columnar cactus, Pilosocereus royenii. Biotropica 38: 635-642. Sabat, A. & J. D. Ackerman. 1996. Fruit set in a deceptive Salzmann, C. C., A. M. Nardella, S. Cozzolino & F. P. deceptive orchids: the signature of pollinator-imposed selection? Ann. Bot. 100: 757-765. Simpson, B. B. & J. L. Neff. 1983. Evolution and diversity in: C. E. Jones and R. J. Little (eds.). Handbook of experimental pollination biology. USA. male and female reproductive success in a deceptively pollinated tropical orchid. Biol. J. Linn. Soc. 151: 405410. effective population size in Lepanthes (Orchidaceae): a case for genetic drift. Biol. J. Linn. Soc. 72: 47-62. Tremblay, R. L., J. D. Ackerman & M.-E. across the selection landscape: annual variation in reproductive characteristics. Philos. Trans. Royal Soc. London, B 365: 491-498. Tremblay, R. L., J. D. Ackerman, J. K. Zimmerman & R. N. Calvo. 2005. Variation in sexual reproduction in orchids Wallace, A. R. 1858. On the tendency of varieties to depart London 3: 46-50. Withner, C. L. 1974. Observations on equitant oncidiums as examples of introgressive hybridization. Pp. 34-45 in: H. H. Szmant & J. Wemple (eds.). First symposium on Chemistry Department, University of Detroit, Detroit, Michigan, USA. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.184 LANKESTERIANA
LANKESTERIANA 11(3): 185. 2011.HOW MANY ORCHID SPECIES IN COSTA RICA? A REVIEW OF THE LATEST DISCOVERIES DIEGO BOGARN email@example.com ABSTRACT. Brassia, Dracula, Lycaste, Polycycnis, Stanhopea, and Trichopilia Sobralia Acianthera aberrans, pendulum, Ornithocephalus montealegrae, and Warmingia zamorana Ecuador. The genus Uleiorchis Uleiorchis ulaei by Ron Liesner, constitutes an interesting new record in Costa Rica. from Costa Rica, has recently been reported from Ecuador. Campylocentrum tenellum, Lepanthes droseroides, Lepanthes mariposa, and Sobralia bouchei from Panama were also lately collected in Costa Rica. Although como: Brassia, Dracula, Lycaste, Polycycnis, Stanhopea, y Trichopilia pero la gran mayora de especies Sobralia Acianthera aberrans, Epidendrum Ornithocephalus montealegrae, y Warmingia zamorana han sido encontradas tanto en Costa Rica como en Ecuador. El gnero Uleiorchis Uleiorchis liesneri descrita de Costa Rica, ha sido registrada recientemente en Ecuador. Campylocentrum tenellum, Lepanthes droseroides, Lepanthes mariposa, y Sobralia bouchei de Panam tambin fueron colectadas recientemente en
Botnico Lankester, asociado con un acceso creciente a la documentacin crtica (tipos y literatura), han sido KEY WORDS : et al., Later, Schlechter (1918), Ames (in Standley, 1937), Central America. The need for an up-to-date checklist Mora-Retana and Garca (1992), Pupulin (2002), and et al. (2007) published a checklist of the orchids of Costa Rica and Mesoamerica. Throughout the history of documentation of the Orchidaceae in Costa Rica, more than three quarters of the species were named by botanists who deposited the material outside the country. Although botanical exploration began almost 150 years ago, sporadically by botanists during the past 15 decades. In the 19th century, Bateman, Lindley, Reichenbach, and Orchid research declined from 1860 to 1900 until Ames named declined dramatically between 1950 and 1960, when less than 30 species were described for Costa Rica, contrasting with the decade of 1920 when nearly there was botanical exploration in those years, the lack of critical materials to identify the species and also the fact that the literature was often published as a management that largely depends on taxonomic and infrastructure, coupled with declining taxonomic has been the lack of adequate information for species strategies such as GTI. With the establishment of Lankester Botanical Garden (LBG) in 1973 and other institutions such as Museo Nacional de Costa Rica there has been an increase in orchid research that has continued until the present. It has been demonstrated (Fig. 1). The mission of LBG has been to eliminate the taxonomic impediment through gathering critical and students. Floristics and botanical exploration in Costa Rica are LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.186 LANKESTERIANA FIGURE 1. Comparison of the number of species of orchids described from Costa Rica at different periods.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 187will not work without an adequate taxonomic basis (Lahaye et al., 2008). The aim of the present paper is to discuss the status of the Orchidaceae in Costa Rica, A review of the latest discoveries After the publication of the most complete treatment Ossenbach et al. (2007) estimated that 1461 orchid species occur in Costa Rica. After Ossenbach et al. (2007), 34 species were described as new to science Rica were recorded for this country. The number now stands at 1519 species. A summary of the latest species known from a single locality or specimen are there with an asterisk. et al. (2007) the species cited, 29 are endemic to Costa Rica. There are 25 species found both in Costa Rica and Panama, and 14 shared with Ecuador. Some disjunctions are recorded, such as Epidendrum stellidifforme and Warmingia zamorana, which are known only from Costa Rica and Ecuador, but their presence in Panama and Colombia is highly probable (Bogarn et al., 2008). Also, there are 16 species widespread from Costa Rica to South America and 10 ranging from Costa Rica to Subtribe Angraecinae: Campylocentrum tenellum specimens of C. tenellum from different localities in Costa Rica are known, but unfortunately in sterile proper recording of this species (Fig. 2a, 3c). An Cordillera de Talamanca. Although C. pachyrrhizum (Rchb.f.) Rolfe is expected to occur in Costa Rica, the material collected indicates that it may correspond to an undescribed species. Schltr. is accepted and considered as distinct from C. fasciola (Lindl.) Cogn. The name C. tyrridion Garay C. dressleri J. Villalobos s.n. (USJ!)) clearly corresponds to Schltr., so C. dressleri is excluded from Costa Rica (Bogarn and Pupulin, 2009). Tribe Gastrodieae: An interesting record of the holomycotrophic orchid genus Uleiorchis is U. ulaei Ron Liesner at MO. The species had been recorded in Subtribe Goodyerinae: Studies published by Ormerod Aspidogyne, A. grayumii Ormerod, known from the Estacin Biolgica Las Cruces in southern Costa Rica near Panama. A new Kreodanthus, K. curvatus Ormerod, is known from Cordillera and also from a collection in the Cordillera de Talamanca. It is compared with the Panamanian K. bugabae Ormerod. Two new species of Microchilus were also recorded M. maasii Ormerod from the M. tessellatus Ormerod, endemic to Costa Rica. The recently described M. valverdei Ormerod is now a synonym of M. calophyllus (Rchb.f.) Ormerod. Also, Platythelys alajuelae Ormerod is endemic to Costa Rica and known from the Cordillera de Tilarn and the Cordillera Central. Subtribe Laeliinae: of Encyclia in Costa Rica yielded a new record: Encyclia gravida (Lindl.) Schltr. Plants of this species considerable in the large genus Epidendrum. The co-workers (2004, 2006, 2008). The following new E. apatotylosum E. astro-selaginellum E. bicuniculatum E. isthmoides E. xnocteburneum hybrid between E. eburneum Rchb.f. and E. nocturnum Jacq.), E. salpichlamys E. steno-
Species Voucher 1. Acianthera aberrans (Luer) Pupulin & Bogarn, Lankesteriana 8(2): 53-55. 2008. Pupulin 4857 JBL 2. Acianthera cabiriae 12(1): 160. 2007. Karremans 1433, CR 3. Acianthera fecunda 12(1): 158. 2007. Bogarn 2650, CR 4. Acianthera hamata 2007. Quesada Chanto s.n., CR 5. Aspidogyne grayumii Grayum 9267, MO 6. Barbosella orbicularis Luer, Selbyana 3(1,2): 10-11, f. 108. 1976. Bogarn 1949, JBL 7. Bensteinia ramonensis Pupulin, Selbyana 28(2): 113. 2008. Bogarn 1923, CR 8. Campylocentrum tenellum Bogarn 5844, JBL 9. Dichaea gomez-lauritoi Gmez-Laurito 8174, CR 10. Dryadella fuchsii Luer, Monogr. Syst. Bot. Missouri Bot. Gard.76: 162-163. 1999. Standley & Valerio 45606, AMES 11. Echinosepala lappiformis M.W.Chase Lindleyana 17(2): 101. 2002. Pleurothallis lappiformis Bot. Missouri Bot. Gard. 15: 38. 1986. Brenesia lappiformis L.O.Williams) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.95: 255. 2004. Echinella lappiformis Lindleyana 16(4): 253. 2001. Dressler 6768, JBL 12. Elleanthus ligularis Dressler 6836B, CR 13. Encyclia gravida (Lindl.) Schltr., Beih. Bot. Centralbl.36(2): 472. 1918. Epidendrum gravidum Pupulin 5377 JBL 14. Epidendrum acroamparoanum (Mexico) 8: t. 802. 2006. Morales 4737, MO 15. Epidendrum atypicum 8: t. 810. 2006. Weber 6081, AMES 16. Epidendrum angustilobum Fawc. & Rendle, J. Bot. 47(556): 124-125. 1909. Epidendrum latifolium 53(3): 392. 1972. Moraga 312, INB 17. Epidendrum apatotylosum Horich sub E. Hgsater 7117, INB 18. Ames & C.Schweinf., Schedul. Orch. 10: 58-59. 1930. Epidendrum powellii 38. 1922. Herrera 3595, CR 19. Epidendrum astroselaginellum 910. 2007. Gmez 23814, MO 20. Epidendrum bicuniculatum 2007. Grayum & G. Herrera 7716, MO 21. Epidendrum buenaventurae 470-471. 1899. Gmez 19984, MO 22. Epidendrum chalcochromum 2006. Hgsater & Mora de Retana 9495, INBLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.188 LANKESTERIANA TABLE
23. Epidendrum cristatum 1798. Epidendrum raniferum Lindl., Gen. Sp. Orchid. Pl.109. 1831. Brade 1281, AMES 24. Epidendrum dolichochlamys (Mexico) 8: t. 829. 2006. Chavarra 661, INB 25. Epidendrum gibbosum L.O.Williams, Ann. Missouri Bot. Gard. 28(4): 420421, pl. 21, 7-10. 1941. Epidendrum acrostigma Icon. Orchid. 3: t. 301. 1999. Grayum 4578, CR 26. Epidendrum haberi 840. 2006. Haber & Cruz 7245, INB 27. Epidendrum hawkesii Horich sub Hgsater 7649, AMO 28. Epidendrum isthmoides 2008. Bello 1516 INB 29. Epidendrum maduroi pl. 352. 1999. Bogarn 1465, JBL 30. 2006. Dressler 6259, AMO 31. Epidendrum nocteburneum 1148. 2008. Weston 57 sub Rodrguez 1042, USJ 32. Epidendrum orthodontum 1999. Karremans 2255, JBL 33. Epidendrum pachytepalum (Mexico) 8: t. 865. 2006. Burger 8253, F 34. Epidendrum pendens L.O.Williams, Ann. Missouri Bot. Gard. 28(4): 421422, pl. 23. 1941. Haber & Cruz 7986, CR 35. Epidendrum philowercklei (Mexico) 8: t. 870. 2006. Fernndez 677, CR 36. Epidendrum platystomoides (Mexico) 8: t. 872. 2006. Hgsater & Mora de Retana 9507 AMO 37. Epidendrum purpurascens Eerste Kl. Kon. Ned. Inst. Wetensch.) 4: 64-65. 1851. Epidendrum glumibracteum Valerio 1017, CR 38. Epidendrum rousseauae Schltr., Beih. Bot. Centralbl., Abt. 2 36(2): 407408. 1918. Todzia 552, CR 39. Epidendrum salpichlamys (Mexico) 8: t. 883. 2006. Grayum & Pam Sleeper 3446, MO 40. Pupulin 6500 JBL 41. Epidendrum serruliferum 44. 1923. B, destroyed; drawing of type, AMES 42. Epidendrum stellidifforme t. 487. 2001. Bogarn 2814, JBL 43. Epidendrum stenoselaginellum 991. 2007. Hgsater 11115, AMO 44. Epidendrum tritropianthum Icon. Orchid. (Mexico) 9(6): t. 998. 2007. Ingram & Ferrell 743 INB 45. Epidendrum villotae Herrera 5151, MOLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 189 Species Voucher
46. Epidendrum vulcanicola Alczar 56, USJ 47. Epidendrum vulgoamparoanum (Mexico) 8: t. 898. 2006. Hgsater 6963, INB 48. Epidendrum zunigae 63. 2008. Bogarn 2680, JBL 49. Gongora boracayanensis 99. 2007. SEL 50. Kefersteinia alata 2004. Carman s.n,, JBL 51. Kefersteinia saccata Pupulin, Willdenowia 38(1): 188. 2008. Pupulin 6549 CR 52. Kreodanthus curvatus Grayum 7481, MO 53. Lepanthes droseroides Luer, Lindleyana 2: 188. 1987. Bogarn 5234, JBL 54. Lepanthes gratiosa 2009. Pupulin 7350 CR 55. Lepanthes machogaffensis 140. 2009. Pupulin 7308 CR 56. Lepanthes mariposa Luer, Phytologia 55: 187. 1984. Bogarn 5034, JBL 57. Lepanthes pelvis Pupulin 7336 CR 58. Lockhartia chocensis 50(83): 19. 1923. Bogarn 2352, JBL 59. Lycaste angelae Oakeley, Lycaste, Ida, Anguloa 27. 2008. 60. Lycaste bruncana Bogarn, Lankesteriana 7(3): 543. 2007. Lycaste crystallina Wubben ex Oakeley, Lycaste Ida, Anguloa 73. 2008. Bogarn 3987, CR 61. Lycaste x daniloi Oakeley, Lycaste, Ida, Anguloa 76. 2008. 62. Lycaste panamanensis Lycaste macrophylla subsp. panamanensis f. 1964. see Oakeley (2008) 63. Maxillaria bolivarensis Whitten 2030 JBL 64. Microchilus maasii Bello 2809 CR 65. Microchilus tessellatus Grayum 8925, MO 66. Ornithidium pendulum (Poepp. & Endl.) Cogn., Fl. Bras.3(6): 92. 1904. Karremans 448, JBL 67. Palmorchis nitida Gmez 26248, USJ 68. Platythelys alajuelae Ingram & Ferrell 680 MO 69. Pleurothallis bogarinii 690. 2007. Pupulin 5285 CR 70. Pleurothallis caudatipetala 22, f. 5-8. 1942. (C. Schweinf.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.95: 259. 2004. Panmorphia caudatipetala (C. Schweinf.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.105: 150. 2006. Luer 12137, MO 71. Pleurothallis duplooyi 50. 2001. (Luer & Sayers) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 260. 2004. Panmorphia duplooyi (Luer & Sayers) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 153. 2006. Bogarn 6955, JBLLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.190 LANKESTERIANA Species Voucher
72. Pleurothallis minutalis Lindl., Fol. Orchid. Pleurothallis 40. 1859. Humboltia minutalis Humboldtia minutalis Panmorphia minutalis (Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.105: 167. 2006. (Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 262. 2004. Anathallis minutalis (Lindl.) Pridgeon & M.W. Chase, Lindleyana 16(4): 249. 2001. W 73. Restrepiella lueri Pupulin & Bogarn, Willdenowia 37(1): 325. 2007. Bogarn 3009, CR 74. Scaphyglottis robusta B.R.Adams, Phytologia 64: 253. 1988. Bogarn 2662, JBL 75. Sobralia aspera Dressler 6783, CR 76. Sobralia blancoi Dressler 6706, CR 77. Sobralia bouchei Ames & C.Schweinf., Schedul. Orchid. 10: 4. 1930. Bogarn 4201, JBL 78. Sobralia pendula Dressler 6746, CR 79. Sobralia rarae-avis Dressler 6793, CR 80. Trichopilia primulina Dressler 7030, CR 81. Uleiorchis ulaei 1958. Wullschlaegelia ulaei Cogn., Fl. Bras. 3(4): 244. 1895. Hammel 11657, MO 82. Warmingia zamorana Warmingia margaritacea B.Johans., Lindleyana 7: 194. 1992. Karremans 1123, JBLselaginellum Costa Rica, and Panama), and E. zunigae Karremans & Bogarn (Fig. 4g, 5d). The following et al. (2006) and recorded by Ossenbach et al. (2007): Epidendrum acroamparoanum E. atypicum E. chalcochromum E. dolicho-chlamys E. haberi E. pachytepalum E. philoE. platystomoides E. salpichlamys E.Santiago, E. tritropianthum and E. vulgoamparoanum E. amparoanum Schltr., which is now treated as a synonym of E. barbeyanum E. pachytepalum, E. and E. vulgoamparoanum are found in both Costa Rica and Panama. New collections regularly: E. buenaventurae E. E. stellidifforme E. villotae Ecuador; E. vulcanicola E. maduroi Epidendrum orthodontum 5b), E. pendens L.O.Williams, and E. rousseauae Schltr. from Panama, which was formerly listed by Pupulin (2002). The following taxonomic changes Epidendrum angustilobum Fawc. & Rendle (=E. latifolium Ames & C.Schweinf. (=E. powellii Schltr.), E. cristatum E. raniferum Lindl.), E. gibbosum L.O.Williams (=E. acrostigma E. purpurascens (=E. glumibracteum Rchb.f.), and the reconsidered E. serruliferum Schltr. Two new species and a new addition will be soon published by Pupulin and Karremans (unpublished). A collection of Scaphyglottis robusta been documented in tropical wet premontane forest of the Cordillera de Talamanca (Fig. 10b). Another specimen was collected in the region of Turrialba in LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 191 Species Voucher
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.192 LANKESTERIANA FIGURE 2. Photographs of: A. Campylocentrum tenellum ; B. Dracula maduroi; C. Lepanthes droseroides Lepanthes gratiosa; E. Lepanthes machogaffensis; F. Lepanthes mariposa; G. Lepanthes pelvis Lycaste bruncana; I. Mormolyca fumea; J. Pleurothallis bogarinii; K. Restrepiella lueri; L. Trichopilia primulina
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 193 FIGURE Barbosella orbicularis; B. Bensteinia ramonensis; C. Campylocentrum tenellum Dichaea gomez-lauritoi
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.194 LANKESTERIANA FIGURE 4. Photographs of: A. Acianthera aberrans; B. Barbosella orbicularis; C. Brenesia lappiformis Epidendrum maduroi; E. ; F. Epidendrum stellidifforme; G. Epidendrum zunigae ; I. ; J. Scaphyglottis robusta; K. Warmingia zamorana (Costa Rica); L. Warmingia zamorana
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 195 FIGURE Echinosepala lappiformis; B. Epidendrum orthodontum; C. Epidendrum stellidifforme Epidendrum zunigae
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.196 LANKESTERIANASubtribe Maxillariinae: The genus Lycaste underwent (2007) and Oakeley (2008). A new species restricted described as Lycaste bruncana Bogarn (Fig. 2h). It is closely allied to L. tricolor Rchb.f. but differs mainly in the twisted column and the shape of the callus and pollinarium. Geographic isolation and molecular two species. Lycaste bruncana the laterdescribed Lycaste crystallina Wubben ex Oakeley. Lycaste brevispatha & Paxton is regarded as a synonym of L. candida Lindl., and, according to Oakeley (2008), L. angelae Oakeley should replace L. brevispatha application of the names Lycaste brevispatha and Lycaste candida Lycaste candida as a nomen nudum. Later, in 1851, J. which Lindley and Paxton transferred to Lycaste in 1853. of L. candida 1863. Oakeley (2008) regarded L. brevispatha as a synonym of L. candida but referring to the name L. candida Lindl. nom. nud. and not to L. candida Lindl. ex Rchb.f. of 1863. The latter is predated by L. brevispatha L. brevispatha L. candida. A careful analysis of the application of the names is needed to clarify the status of the species of this group. A natural hybrid Lycaste x daniloi Oakeley is also reported as a cross between L. angelae and L. candida. Lycaste angelae rubra Oakeley was therefore it should regarded as an illegitimate name according to the the International Code of Botanical Nomenclature type must be a specimen after 1 January 2007. The nomenclature of the Lycaste macrophylla (Poepp. & Endl.) Lindl. complex, formerly represented in Costa Rica by three subspecies (L. macrophylla subsp. desboisiana (Cogn.) Fowlie, L. macrophylla subsp. FIGURE 6. Photographs of: A. Acianthera hamata; B. x Bensteinia ramonensis; C. Elleanthus ligularis Epidendrum isthmioides; E. Kefersteinia saccata; F. Ornithidium pendulum of LBG.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 197puntarenasensis Fowlie, and L. macrophylla subsp. Fowlie), also changed in Oakeley (2008). to Costa Rica the following species: (Fowlie) Oakeley, L. panamanensis Fowlie ex Oakeley L. puntarenasensis (Fowlie) Oakeley, and L. measuresiana (B.S.Williams) Oakeley. Lycaste macrophylla subsp. desboisiana was treated as synonym of L. measuresiana (Oakeley, so that the correct names are Lycaste puntarenasensis (Fowlie) Archila, (Fowlie) Archila, and Lycaste panamanensis (Fowlie) Archila. from (Blanco et al., 2007). bolivarensis distributed throughout South America has been recorded based on two collections along the Caribbean lowlands (Fig. 4I, 7b); the recent nomenclatural changes did not affect this name (Blanco 2008). Ornithidium pendulum (Poepp. & Endl.) Cogn. (= (Poepp. & Endl.) C.Schweinf.), a widespread species ranging from Guatemala, Nicaragua, Panama, and South America, is misapplied name et al. (2008). described from Costa Rica, has recently been reported from Ecuador (M. Blanco, personal communication). A new species of Mormolyca, allied to what we call the alliance, was described by Bogarn and Pupulin (2010) (Fig. 2i). Tribe Neottieae: Palmorchis nitida of southern Costa Rica by Bainbridge and Aguilar (2008). Subtribe Oncidiinae: A collection of similar to L. micrantha Rchb.f. has been reported as the Colombian collections from the tropical wet forest-premontane belt transition on the Caribbean watershed of the Cordillera de Tilarn range northwestern Costa Rica (Fig. 4h, 7a). Studies in Trichopilia T. primulina locality of this species. The only record of Warmingia in Costa Rica, W. margaritacea B. Johans., is now considered a synonym of the Ecuadorian W. zamorana et al., 2007; Fig. 4k, 4l, 11c, 11d). Subtribe Pleurothallidinae: This is one of the largest groups represented in Costa Rica. After the studies by Pupulin (2002), Pupulin and Bogarn (2007), and Pupulin et al. species of Acianthera A. cabiriae A. fecunda these from the Caribbean lowlands of the Turrialba region, and A. hamata Pupulin & G.A.Rojas, from an unknown locality (Pupulin et al., 2007; Fig. 8 d). A collection from southern Cartago corresponds to the A. hamata. All the new species are endemic to Costa Rica (Fig. 6a). A new record, A. aberrans (Luer) Pupulin & Bogarn has been found along the Caribbean lowlands in tropical rain forest of the Sarapiqu region (Fig. 4a, 8a). This species was described from Veraguas, Panama. The Panamanian Barbosella orbicularis Luer, along the Caribbean watershed of the Cordillera Central (Fig. 3a, 4b). A specimen of Dracula maduroi Luer was documented in a recent expedition to Volcn Cacho Negro in the Braulio Carrillo National Park in the Cordillera Central (Fig. 2b). A specimen of Dryadella fuchsii Costa Rica by Luer (2005) based on collections by P. C. Standley and J. Valerio around the Tilarn area in the northwestern Costa Rica. The species had not been listed for Costa Rica by Luer (2003) or Ossenbach et al. (2007). The Nicaraguan Echinosepala lappiformis (=Brenesia lappiformis Luer was recorded based on two collections from the 5a). Another specimen was collected near Gupiles, Studies on Lepanthes yielded many new records and species in last few years. Three species L. gratiosa L.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.198 LANKESTERIANA FIGURE ; B. ; C. Ornithidium pendulum Pleurothallis bogarinii
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 199 FIGURE Acianthera aberrans; B. Acianthera cabiriae ; C. Acianthera fecunda Acianthera hamata.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.200 LANKESTERIANA FIGURE Kefersteinia saccata; B. Lepanthes mariposa; C. Lepanthes machogaffensis Lepanthes pelvis.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 201 FIGURE Restrepiella lueri; B. Scaphyglottis robusta; C. Sobralia aspera Sobralia blancoi
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.202 LANKESTERIANA FIGURE Sobralia pendula; B. Trichopilia primulina; C. Warmingia zamorana Warmingia zamorana
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 203machogaffensis and L. pelvis from the Cordillera de Talamanca were recorded from a single path in Tapant National Park. New been registered for L. pelvis and L. machogaffensis. Also, L. droseroides Luer (Fig. 2c), L. mariposa Luer (Fig. 2f, 9b), and Luer (of which it collected in Costa Rica. Unpublished manuscripts unpublished; Pupulin and Bogarn, unpublished). (Luer) Luer was cited by Pupulin (2002) and Luer (2003) based on a plant that its origin is doubtful in all the collection details near Cerro de La Muerte, along the Panamerican of this species in Costa Rica. A new Pleurothallis, P. bogarinii P. saphipetala Luer and restricted to the Caribbean lowlands, has been registered based on two collections (Fig. 2j, 7d). Restrepiella has a second species: Restrepiella lueri Pupulin & Bogarn (Fig. 2k, 10a). It differs from Restrepiella ophiocephala hirsute at apex and ciliate along the margins, twice as long as the column and as long as the petals, the column without a foot and the free lateral sepals. Two additional records added by Luer (2006) are: Pleurothallis caudatipetala C.Schweinf. (= caudatipetala (C.Schweinf.) Luer, =Panmorphia caudatipetala from Peru and Ecuador and collected in Costa Rica around Cascajal, northern San Jos; and P. minutalis Lindl. (Panmorphia minutalis (Lindl.) Luer, = (Lindl.) Luer, = Anathallis minutalis (Lindl.) Pridgeon & M.W. Chase), known from Mexico and Guatemala, and based on a collection of A.R. Endrs (, W). A specimen of Pleurothallis duplooyi Luer & Sayers (= duplooyi (Luer & Sayers) Luer, = Panmorphia duplooyi (Luer & Sayers) Luer), formerly considered watershed of the Cordillera de Talamanca near Pejibaye peduncle. The specimen collected in Costa Rica shows an elliptic dorsal sepal rather than oblong, and basally the drawing published by Luer (2006). The plant was at JBL. There is currently a project on the taxonomy and phylogeny of the Pleurothallidinae in Costa Rica the future. Tribe Sobralieae: Sobralia new species. Sobralia aspera (Fig. 9c), S. blancoi S. pendula S. raraeavis the Panamanian S. bouchei Ames & C.Schweinf, was found in Turrialba on the Caribbean lowlands. Three other new species of Sobralia are awaiting publication Elleanthus, there is a new species, E. ligularis collected in Colombia, Costa Rica, Cuba, and Panama, but the species had remained unnamed in the absence E. graminifolius (Barb.Rodr.) Ljtnant, but the plants are more robust rather than obtriangular, with a short base. Subtribe Stanhopeinae: A new Gongora, Gongora boracayanensis the name Gongora quinquenervis Gongora from Costa Rica (Jenny et al., 2007).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.204 LANKESTERIANASubtribe Zygopetalinae: This subtribe has been well (2007). x Bensteinia ramonensis Pupulin, a natural bigeneric hybrid (Benzingia reichenbachiana (Schltr.) x Dichaea of Costa Rica (Pupulin, 2007), a new species, Dichaea gomez-lauritoi Pupulin, was described from a specimen collected in Gupiles along the Caribbean plains of Limn (Fig. 3d). Four name changes are noted: Dichaea amparoana Schltr. (described D. Dichaea standleyi Ames was placed as a synonym of D. acroblephrara Schltr. Dichaea schlechteri D. similis D. cryptarrhena Dichaea ciliolata Rolfe was reduced to a synonym of D. hystricina Rchb.f. based Kefersteinia (Pupulin, 2001): Kefersteinia saccata Pupulin from the Caribbean lowlands (Pupulin and Merino, 2008; Fig. 6e, 9a), and a new record, the Panamanian K. alata, side of the border with Panama (Pupulin, personal communication). ACKNOWLEDGEMENTS Telecommunications (MINAET) and its National System permits under which wild species treated here were Karremans, and Franco Pupulin for their continuous management of the collections at JBL. The present work is of Costa Rica. LITERATURE CITED Archila, F. 2002. Nuevas combinaciones en el gnero Lycaste (Orchidaceae). Rev. Guatemalensis 5: 58-62. Bainbridge, C. V. & R. Aguilar. 2008. A new addition to Palmorchis nitida (Orchidaceae: Neottieae) is documented from the Osa Peninsula. Lankesteriana 8: 1-4. Blanco, M. A., G. Carnevali, M. Whitten, R. B. Singer, L. Endara. 2007. Generic realignments in Maxillariinae (Orchidaceae). Lankesteriana 7: 515-537. Blanco, M. A., G. Carnevali, D. Bogarn & R. B. Singer. 2008. Further disentangling of a taxonomic puzzle: Maxillaria ramosa, Ornithidium pendulum, and a new species, O. elianae (Orchidaceae). Harvard Pap. Bot. 13: 137-154. Bogarn, D. 2007. A new Lycaste (Orchidaceae: Maxillarieae) from Costa Rica. Lankesteriana 7: 543-549. Bogarn, D., A. Karremans & F. Pupulin. 2008. New species and records of Orchidaceae from Costa Rica. Lankesteriana 8: 53-74. Bogarn, D. and Pupulin, F. 2009. The genus Campylocentrum (Orchidaceae: Angraecinae) in Costa Rica: some critical questions and a few answers. Pp. 3245 in: A. M. Pridgeon and J. P. Surez (eds.). Proceedings Bogarn, D. & F. Pupulin. 2010. Two new species of Mormolyca from Costa Rica and Panama. Orch. Digest 74: 43-47. Dressler, R. L. 1993. Field guide to the orchids of Costa Rica and Panama. Cornell University Press, Ithaca, New York, USA. Dressler, R. L. 2003. Orchidaceae. Pp. 1-595 in: B. E. Hammel, M. H. Grayum, C. Herrera & N. Zamora (eds.). Manual de plantas de Costa Rica Volumen III, monocotiledneas (Orchidaceae-Zingiberaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 93. Dressler, R. L. & D. Bogarn. 2007. Elleanthus ligularis, a name for a relatively common new species of Elleanthus sect. Chloidelyna Lankesteriana 7: 539-542. Dressler, R. L. & D. Bogarn. 2009. Der Trichopilia tortilis komplex (Orchidaceae: Oncidiinae) mit einer schwierig zu bestimmenden neuen art. The Trichopilia tortilis complex (Orchidaceae: Oncidiinae) with an elusive new species. Orchideen J. 2: 56-65. Hgsater, E., L. Snchez-Saldaa & J. Garca-Cruz. 2004. Icones Orchidacearum, Fasc. 7. The Genus Epidendrum. Part 4. Instituto Chinoin, A.C., Mxico. Hgsater, E., L. Snchez-Saldaa & J. Garca-Cruz. 2006. Icones Orchidacearum, Fasc. 8. The Genus Epidendrum. Part 5. Instituto Chinoin, A.C., Mxico. Hgsater, E. & L. Snchez-Saldaa. 2008. Icones Orchidacearum, Fasc. 11. The Genus Epidendrum. Part 7. Instituto Chinoin, A.C., Mxico. Hemsley, W. B. 1883. Orchideae. Pp. 197-308 in: F. D. Goldman & O. Salvin (eds.). Biologia Centrali-
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BOGARN 205Americana. R. H. Porter, London, UK. Jenny, R., S. Dlstrom & W. H. Higgins2007. An old species Gongora boracayanensis. Selbyana 28: 99-102. Lindley, J. 1851. Lycaste leucantha Paxtons Flower Gard. 2: 37. Lahaye, R., M. van der Bank, D. Bogarn, J. Warner, F. Pupulin, G. Gigot, O. Maurin, S. Duthoit, T. Barraclough biodiversity hotspots. Proc. Nat. Ac. Sc. USA 105: 29232928. Luer, C. A. 1987. New Lepanthes species from Costa Rica and Panama. Lindleyana 2: 185-217. Luer, C. A. 2003. Pleurothallis Pp. 386-452 in: B. E. Hammel, M. H. Grayum, C. Herrera & N. Zamora (eds.). Manual de plantas de Costa Rica Volumen III, monocotiledneas (Orchidaceae-Zingiberaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 93. Luer, C. A. 2005. Icones Pleurothallidinarum XXVII. Dryadella and Acronia section Macrophyllae Fasciculatae Monogr. Syst. Bot. Missouri Bot. Gard. 103: 1-311. Luer, C. A. 2006. Icones Pleurothallidinarum XXVIII. Reconsideration of Masdevallia and the systematics of Specklinia and vegetatively similar genera (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 105: 21-244. Mora-Retana, D. E. & J. B. Castro. 1992. Lista actualizada de las orqudeas de Costa Rica (Orchidaceae). Brenesia 37: 79-124. Oakeley, H. F. 2008. Lycaste, Ida and Anguloa: the essential guide. The author, Beckenham, Kent, UK. Ossenbach, C., F. Pupulin & R. L. Dressler. 2007. Orqudeas del istmo centroamericano: catlogo y estado de conservacin/Checklist and conservation status: Orchids of the Central American isthmus. Editorial 25 de Mayo, Sabanilla de Montes de Oca, San Jos, Costa Rica. Ormerod, P. 2007. Studies of Neotropical Goodyerinae (Orchidaceae). Harvard Pap. Bot. 9: 391-423. Ormerod, P. 2008. Studies of Neotropical Goodyerinae (Orchidaceae). Harvard Pap. Bot. 13: 55-87. Pupulin, F. 2001. Contributions to a reassessment of Costa Rican Zygopetalinae (Orchidaceae). The genus Kefersteinia Rchb.f. Ann. Naturhisto. Mus. Wien 103B: 525-555. Pupulin, F. 2002. Catlogo revisado y anotado de las Orchidaceae de Costa Rica. Lankesteriana 4: 1-88. Pupulin, F. 2005. Ciliate dichaeas; Dichaea hystricina and Dichaea ciliolata : two species in one and an interesting variation. Orchids 74: 678-683. Pupulin, F. 2007. Contributions toward a reassessment of Costa Rican Zygopetalinae (Orchidaceae). 3. A systematic revision of Dichaea in Costa Rica. Harvard Pap. Bot. 12: 15-153. Pupulin, F. & D. Bogarn. 2004. Two new species of Lepanthes (Orchidaceae: Pleurothallidinae) from Costa Rica. Kew Bulletin 59: 559-563. Pupulin, F. & D. Bogarn. 2007. A second new species in the genus Restrepiella (Orchidaceae:Pleurothallidinae). Willdenowia 37: 323-329. Pupulin, F., D. Bogarn & D. Jimnez. 2009. New species and records in Mesoamerican Lepanthes. Orch. Digest 73: 136-145. Pupulin, F. & G. Merino. 2008. Two new species of Kefersteinia (Orchidaceae: Zygopetalinae). Willdenowia 38: 1-7. species of Acianthera (Orchidaceae: Pleurothallidinae) from Costa Rica. Harvard Pap. Bot. 11: 155-162. Reichenbach, H. G. 1866. Beitrge zu einer orchideenkunde Central-Amerikas. Hamburg, Germany. Schlechter, R. 1918. Kritische aufzhlung der bisher aus Zentral-Amerika bekanntgewordenen. Orchidaceen, Beihefte zum Botanischen Centralblatt Verlag von C. Heinrich, Dresden, Germany. Standley, P. C. 1937. Flora of Costa Rica (Part 1). Publications of Field Museum of Natural History, Botanical Series 18: 1-63. Williams, L. O. 1956. An enumeration of the Orchidaceae of Central America, British Honduras and Panama. Ceiba 5: 1-2.
LANKESTERIANA 11(3): 207. 2011.MATING SYSTEMS IN THE PLEUROTHALLIDINAE (ORCHIDACEAE): EVOLUTIONARY AND SYSTEMATIC IMPLICATIONS EDUARDO LEITE BORBA ARIANE RAQUEL BARBOSA, MARCOS CABRAL DE MELO, SAMUEL LOUREIRO GONTIJO & HENRIQUE ORNELLAS DE OLIVEIRA Corresponding author: firstname.lastname@example.org ABSTR A CT. pollinations of species in the major genera representing all major lineages of Pleurothallidinae in order to determine Octomeria, a genus with moment, all but one of the species studied in selected large genera of the major lineages (Acianthera, Anathallis, Masdevallia, Octomeria, and Stelis) are self-incompatible. The species studied may possess complete, strong or partial self-incompatibility. We found two different sites where self-incompatibility reactions occur, the stigma and the stylar channel, and both sites were not found in the same genus except for Anathallis. In Anathallis, the two groups that differ morphologically (formerly Pleurothallis subgen. sect. Muscosae and sect. Acuminatae) exhibit different sites of reaction. Flowers of Octomeria species produce nectar and are pollinated in Acianthera s.s. condition. In addition to the morphological synapomorphy uniting the members of the former circumscription biological synapomorphies in the group self-incompatibility and myophily. Based on these differences, we self-incompatible, myophilous clade and the other of the small, self-compatible, ornithophilous clade. RESUMEN Octomeria, un gnero con caractersticas tpicas de plantas todas excepto una, de las especies estudiadas de los gneros seleccionados (Acianthera, Anathallis, Masdevallia, and Stelis) son auto-incompatibles, pudiendo ser fuerte o parcialmente auto-incompatibles. Se encontr dos sitos donde las reacciones de auto-incompatibilidad podran ocurrir, siendo stas el estigma y el canal del estilo. Los dos sitios de auto-incompatibilidad no fueron encontrados dentro de las especies del mismo gnero, a excepcin de Anathallis Pleurothallis subgen. sect. Muscosae and sect. Acuminatae) presentan diferentes sitios de autoOctomeria especies de Acianthera
. KEY WORDS : LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.208 LANKESTERIANA orchids is mostly attributed to the striking adaptations of these plants to attract pollinators, which range from insects to birds. The morphological adaptations considered largely responsible for the great numbers pollination barriers, which may be morphological, Cocucci, 1999). Apparently this mating system helps allogamy by pre-pollination barriers are found scattered throughout the family, such as agamospermy, self-pollination, and self-incompatibility (Tremblay et al., 2005). In contrast to agamospermy and autonomous self-pollination, which are more common in Orchidoideae, self-incompatibility is more frequent in Epidendroideae, mainly in Cymbidieae, Vandeae, 1982; Tremblay et al., 2005). Genetic barriers (i.e., (Agnew, 1986; Johansen, 1990; Christensen, 1992; associated with species pollinated by insects that may promote autogamy or geitonogamy (Christensen, 1992; Pedersen, 1995, Borba & Semir, 1998, 1999, 2001; Singer & Cocucci, 1999; Borba et al., 2001a). The main pollinators of Orchidaceae are members 1966; Proctor et al. exploring more complex mechanisms of pollination, such as relationships with male euglossine bees, pseudocopulation mechanisms, and more species(Christensen, 1994; Proctor et al. mainly in the Pantropical Bulbophyllum and the Neotropical Pleurothallidinae (Epidendreae; Borba & Semir, 1998, 2001; Tan et al., 2002, 2006). These two unrelated taxa are the main groups of myophilous orchids (Christensen, 1994). Myophily also occurs in many other unrelated lineages within the family, indicating that pollination by dipterans is a 1993; Christensen, 1994). Pleurothallidinae comprise about 4100 species in 37 genera, encompassing 20% of the species of the family, and is the largest orchid group pollinated by by the absence of pseudobulbs and the presence of An exception is seen in the clade composed by three small genera from Central America (Dilomilis, and Tomzanonia), currently included in the subtribe (Pridgeon & Chase, 2001; Pridgeon et al., 2001, 2005; (Fig. 1). In recent molecular phylogenetic
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae209 FIGURE matK/trnL/ et al. below branches are equally weighted bootstrap percentages >50%. Letters indicate the main clades according to Pridgeon et al. Laeliinae. Outgroups are represented by Arpophyllum giganteum and Isochilus amparoanus. Reproduced from Pridgeon & Chase (2001) with permission.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.210 LANKESTERIANAanalyses, in spite of being a monophyletic and easily enlarged to include that small clade (a total of only eight species) of bird-pollinated, self-compatible (Pridgeon et al., 2001, 2005). In the combined analysis (matK and trnL-F recent phylogenetic analysis, the clade consisting of Octomeria and Brachionidium is the sister group to the rest of the members in the older delimitation of the subtribe (sensu trnL-F and ITS separate analyses, Octomeria alone is the sister group of the remaining Pleurothallidinae (sensu Luer, 1986). The whole larger subtribe is sister to Laeliinae. Laeliinae are primarily self-compatible birdand lepidoptera-pollinated groups and a few selfBerg et al., 2005, 2009). morphology, odor, and coloration typical of plants presence of mobile appendages, unpleasant smell, and dull purplish and yellow colors with spots and almost all species of Pleurothallidinae are pollinated authors to be a plesiomorphic syndrome in the subtribe of at least two of the three genera of the clade recently included in the subtribe are apparently pollinated by hummingbirds (Ackerman, 1995; Pridgeon et al., 2005). Another important exception is the genus Octomeria, which is represented by approximately 150 species with a Neotropical distribution, concentrated mainly in northern South America and southeastern the genus, and it is typical in morphology, odor, and coloration typical of plants pollinated by bees (Fig. 2Fet al., 2004). These facts led us to question when myophily arose in the subtribe before becoming a widely occurring characteristic in the group. Pleurothallidinae: one species of Stelis (Christensen, 1992), three of Lepanthes (Tremblay et al., 2005), and Acianthera (Borba et al. included in Pleurothallis subgen. Acianthera; Pridgeon and Chase, 2001; Borba, 2003), which are of Acianthera studied by Borba et al. (2001a) had reaction-site and pollen-tube morphology similar gametophytic self-incompatibility (de Nettancourt, et al., 1996; Richards, 1996). They found some features that may indicate a distinct incompatibility system, but only diallelic crossing and embryology studies can or sporophytic) is shown by those species (Richards, 1996; Lipow & Wyatt, 2000). Dilomilis is self-compatible (Ackerman, 1995). This is a plesiomorphic characteristic in Laeliinae (Borba Epidendrum species (Adams & Goss, 1976; Pansarin, 2003). The distribution of self-incompatibility in remaining Epidendreae indicates that it is also a are typically self-compatible, we could expect that Acianthera species studied by Borba et al. (2001b) et al., 1989; Scacchi et al., 1990; Corrias et al., 1991; Klier et al., 1991; Case, 1994) and near the maximum known et al. (2001a) suggested that both self-incompatibility and inbreeding depression are responsible for the Bulbophyllum et al., 2007).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae211 FIGURE Acianthera limae; B. A. saurocephala ; C. Anathallis microphyta A. sclerophylla ; E. Masdevallia infracta ; F. Octomeria campos-portoi; G. O. crassifolia, with pollinator Bradysia O. ; I. O. wawrae; J. ; K. Stelis aff. peliochyla ; L. Stelis sp.; M. S. aff. hypnicola ; N. Acianthera prolifera Acianthera prolifera; note the least amount of seeds in the self-pollinated fruit. Facing this scenario found so far in Pleurothallidinae, subtribe in response to a selection pressure caused by the change from a group that promotes crossin the group, populations were able to maintain myophily) with self-incompatibility or myophilous
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.212 LANKESTERIANA TABLE et al.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae213 TABLE et al., 2001; see O). pollinations of species in the major genera representing all of the major lineages of Pleurothallidinae in order to determine: 1) the extent of occurrence of selfincompatibility in the subtribe; 2) in which group it has Octomeria in order to determine if myophily arose at the base of the clade that corresponds to the older circumscription of the subtribe or if it arose later in phylogeny. Materials and methods Mating systems We sampled 22 species in eight et al. (2001; Fig. 1, Table 1): Acianthera (six species), Anathallis (four spp.), Masdevallia (one), (one), Octomeria (four), (two), Stelis (three), and Zootrophion (one). It was not possible to sample Phloeophila species, the only genus of this lineage. Studies with the species sampled (clade B) are still in progress and are not presented here. Thus,
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.214 LANKESTERIANAwe present the results of six of the eight main clades of Pleurothallidinae. Vouchers were deposited in the The populations studied were located in areas of campo rupestre (Minas Gerais and Rio de Janeiro states) and occurring as epiphytes in semideciduous, gallery, and Atlantic rain forests or as rupicolous on rock outcrops. and maintained for a minimum of six months in a Gerais. Three types of pollination experiments were in each population. All populations were subjected to self-pollination and intrapopulation cross-pollination pollinations per treatment. Interpopulation crosses were performed in some species with a multipopulation per population and at least two populations per species. Sometimes the numbers were lower, but in some cases they were as high as 160 pollinations per treatment, species. Additionally, some marked emasculated or not the possible occurrence of diplosporic agamospermy pollination experiments were checked on a daily basis Additional selfand cross-pollinations were 60 C for approximately 25 min. (or 50 min. in the case of fruits). The material was then washed in distilled water and stained with Aniline Blue for examination 1959). The fruits were collected at the start of their 300 seeds from each fruit were examined by optical on a morphological basis only, according to the seeds with no embryo or a rudimentary embryo were et al., 2001a). Floral biology of Octomeria speciesField O. crassifolia and in the Serra do Caraa Mountains (municipality of Catas Altas, Minas Gerais were made on January and February 2007 from 07:00 to O. crassifolia were made on January 2007, between 06:00 and 12:00, totaling 63 only a summary description of the pollination of the species with the identity of pollinators, because a species is being published elsewhere (Barbosa et al., 2009). Results Flowers that did not produce fruits abscised within 5-10 days, except for Acianthera prolifera and Anathallis microphyta Anathallis heterophylla, and Acianthera, Octomeria, Stelis species), 110-130 days (other Anathallis species) or 150 days (Anathallis sclerophylla and Masdevallia infracta Fruit set in experimental intra-population crossfrom 5 to 88% (Table 2). Fruit set in inter-population cross-pollinations was similar to that found in intrapopulation crossings, the latter being usually slightly lower or more rarely higher (e.g., Acianthera limae). Except for one species, fruit set in self-pollinations was always lower than in cross-pollination experiments,
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae215 FIGURE Anathallis rubens Octomeria crassifolia Stelis aff. peliochyla , after Masdevallia infracta with normal pollen tubes reaching the base of the column; J. Pollen tubes in a mature Masdevallia infracta Acianthera saurocephala two species). Ten species did not set any fruits in selfpollinations, and fruit set in this treatment was higher than in selfpollinated fruits. The latter also had fewer seeds, and sometimes they failed to open, mainly in Acianthera for Zootrophion atropurpureum, fruit set in both selfand cross-pollinations was nearly 55%. In this species, and seed content in both treatments. Although Masdevallia infracta number of self-pollinated fruits (n=9) had 67%. The self-pollination (n=2, 2, and 9), but they did in crosspollination (one of them with 100% fruit set in four cross-pollinations), indicating that they are not sterile. Bidirectional crossings between one pair of self-
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.216 LANKESTERIANA was the pollen donor (n=4). Another pair of similar always high in cross-pollinated fruits, usually greater than 80% (except for 74%, and O. praestans ual fruits was possibly as low as 7% than 70% (except for Masdevallia infracta, 73%), and always lower than in cross-pollinated capsules (including Z. atropurpureum as 100% but more frequently lower than 20%. We examined of any species. Pollen germination and pollen-tube growthIn experimental cross-pollination, germination of pollen day after pollination, depending on the species. In this pollination treatment, practically all pollen grains germinated, and the pollen tubes demonstrated uniform growth, with callose plugs deposited regularly along for Anathallis microphyta and Acianthera species, for which it occurred on the 11th day. Penetration of the in cases of cross-pollination. We found two general patterns for pollen and pollenMasdevallia infracta and all Octomeria, Stelis, and Anathallis (except A. microphyta) species, pollen grains in the majority that did initiate germination demonstrated irregular growth and acquired a twisted appearance when they reached the initial portion of the style with irregular deposition of callose on the pollen tube walls and at the extremity of the tubes rarely reac Acianthera species and Anathallis microphyta, pollen-tube growth occurred from that point onward, pollen tubes began to take on abnormal characteristics that had reached the base of Acianthera saurocephala the pollen tubes assuming abnormal appearance as Masdevallia infracta that set fruit in self-pollinations, pollen tubes Pollen tubes in fruits with a high percentage of fruits with a high percentage of empty seeds (lacking embryos) usually showed normal pollen tube growth only in the column; they became irregular when pollinations, mainly in Acianthera species, which showed higher fruit set in this treatment than the other species. Floral biology of Octomeria speciesOctomeria crassifolia and females in similar proportions of four species of Bradysia early in the morning, between 06:00 and 08:00. Pollinators of O. crassifolia a single species of Pseudosciara O. crassifolia
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae217pollinators of O. crassifolia but no pollinarium was The pollinators of the two species demonstrated to the labellum, which shifts slightly downward. labellum, feeding on the nectar produced on the disk. population of O. crassifolia O. Discussion Absence of fruit set in unpollinated or emasculated fruits by autonomous self-pollination or agamospermy; thus, a pollination is required for the formation of fruits in these species. Absence or low fruit set in cross-pollinations indicates occurrence of complete or strong self-incompatibility in the majority of the species studied (see Table 2). Complete or strong self-incompatibility was also found in the majority of Pleurothallidinae species studied so far (Christensen, 1992; Borba et al., 2001a; Tremblay et al., 2005; see Table 2). study in Acianthera (Borba et al., 2001a), fruit set by Borba et al. (2001a), strict self-incompatibility and selfcompatibility are extremes of a continuum between which there is often no clear-cut difference, The distinction between the two conditions is arbitrary, indices to determine whether a particular species is self-compatible or self-incompatible (e.g., Bawa species in this study (e.g., Acianthera prolifera) could be considered self-incompatible or self-compatible depending on which method is employed. Some fruit set, which we think is improper, because seed If that practice is used for these species, they would be considered clearly self-incompatible on account timing of the reaction in the stylar canal or absence of pollen germination lead us to suggest the occurrence of partial self-incompatibility in these species. Zootrophion atropurpureum can be clearly considered self-compatible, in spite of the reduced seed to crosspollination. But Masdevallia infracta, which could be considered self-compatible if based simply on total fruit set, probably has a more complex mating population of Masdevallia infracta exhibit strong selfincompatibility (with the reaction site on the stigma because of little or no pollen germination), and half the as we could see by the bundle of normal pollen tubes between self-compatible and self-incompatible responsible for the stylar self-incompatibility factor. Thus, the species, or at least this population, may L., with incompatible population (Lipow & Wyatt, 2000). This may also explain the occurrence of some fruit set in strong or partial self-incompatibility. Unfortunately, the Z. atropurpureum was low, and so we cannot know whether the same occurs in this species/ population or if it is in fact self-compatible. We will try to increase the number of experimental pollinations and The species showed two different selfincompatibility reactions, one typical of gametophytic self-incompatibility (pollen tubes becoming irregular
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.218 LANKESTERIANA of sporophytic self-incompatibility (absence of pollen grain germination; Richards, 1996). In addition to one Anathallis species, the reaction typical of gametophytic Acianthera species. There are currently no published although they are fundamental to understanding control of self-incompatibility in the group. We need more studies to determine if this picture represents either two incompatibility systems (gametophytic and sporophytic) or two sites of the same reaction. Intermediate situations, as found in species such as Acianthera saurocephala and Stelis aff. hypnicola, indicate that the latter scenario is more plausible. All Anathallis species showed strong selfincompatibility, but we found the two different sites where self-incompatibility reactions occur, the stylar channel and the stigma, which are related to Pleurothallis subgen. sect. Muscosae and P. subgen. Acuminatia (Luer, 1986, 1999). This differences in morphology, the new circumscription groups were combined to form the core of the genus Anathallis (Pridgeon & Chase, 2001). This is the only genus in which both sites of incompatibility were found. In this same clade, there is a report of strong self-incompatibility in three Lepanthes species by Tremblay et al. (2005). In Anathallis microphyta, formerly included in Pleurothallis subgen. sect. Muscosae and recently transferred by Luer (2006) to Panmorphia, germination of the pollen incompatibility reaction occurs when the pollen tubes reach the base of the column. On the other hand, in the three species formerly belonging to Pleurothallis subgen. Acuminatia, the pollen grains fail to germinate or pollen germination is low and the pollen tubes do order to reach a better understanding of the distinct incompatibility mechanisms in the genus, additional studies examining the mating systems and molecular phylogenetic analysis are necessary, both using larger samples of species belonging to the two morphological groups. Experiments in progress by our group indicate that self-incompatibility can also be assigned to other species of Pleurothallidinae, such as Octomeria campos-portoi, O. diaphana, O. wawrae, M. punctatus, and additional Acianthera and Occurrence of self-incompatibility in species is particularly important to our study, since they represent a clade not sampled in this study (see Fig. 1). Our results indicate that self-incompatibility is myophilous clade of the subtribe, in spite of the occurrence of self-compatibility in Zootrophion, Octomeria species ( and O. crassifolia et al., unpublished), and Acianthera incompatibility and inbreeding depression apparently et al., 2001a, 2001b). The phylogenetic position of Octomeria indicates that self-incompatibility and myophily arose at group of the subtribe, corresponding to the older 1993). As such, self-incompatibility and myophily may be considered as biological synapomorphies of this large myophilous clade. The concomitant occurrence of myophily and self-incompatibility in all of the clades studied indicates two hypothetical routes for the
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae219arisen as a result of selection pressures that arose by the change of pollinators, from an insect group with time, our results cannot distinguish which of the two Sapromyophily is apparently widely distributed and represents the predominant pollination mechanism in 1994). Some characteristics of Octomeria a yellow perianth, nectar guides on the calli, nectar reward at the base of the lip, and agreeable citrony odor are found in myophilous species but are also common in mellitophilous species. Because of these and other purple lip, Octomeria we suggest that myophily sensu stricto (pollination a plesiomorphic characteristic in the myophilous clade of the subtribe in spite of the most common occurrence of sapromyophily (pollination by female is not uncommon in orchids, and the application of these cases (e.g. Pansarin, 2008). In the older circumscription of the subtribe, the considered an important diagnostic characteristic for the of the small clade containing the genera Dilomilis, and Tomzanonia this characteristic), no morphological synapomorphy et al., 2001). For this reason, and the occurrence of self-compatibility and ornithophily in some species of this clade, we suggest that the older circumscription of the subtribe should be maintained with self-incompatibility and myophily as biological synapomorphies and the articulation as a morphological synapomorphy for Pleurothallidinae. As such, the establishment of a subtribe comprising the genera Dilomilis, and Tomzanonia should not be the main argument for its inclusion in the subtribe (see Pridgeon et al., 2001). As these two groups are sister to each other, such taxonomic rearrangement is still supported by the results found by Pridgeon et al. (2001). ACKNOWLEDGMENTS. in the RPPN Serra do Caraa, Rubens Custdio da Mota for information on the location of plants and all the staff of the Setor de Biossistemtica e Sistemtica Molecular in the Laboratrio de Sistemtica Vegetal, UFMG, and was supported by grants from the Conselho Nacional de from the Pr-Reitoria de Pesquisa/UFMG. Eduardo Borba LITERATURE CITED and the Virgin Islands. Mem. New York Bot. Gard. 73: 1-203. Adams, R. M. & G. J. Goss. 1976. The reproductive biology of the epiphytic orchids of Florida III. Epidendrum anceps Jacq. Amer. Orch. Sco. Bull. 45: 488-492. Agnew, J. D. 1986. Self-compatibility/incompatibility in some orchids of the subfamily Vandoideae. Pl. Breed. 97: 183-186. Azevedo, M. T. A., E. L. Borba, J. Semir & V. N. Solferini. 2007. High genetic variability in Neotropical myophilous orchids. Bot. J. Linn. Soc. 153: 33-40. Barbosa, A. R., M. C. Melo & E. L. Borba. 2009. Selfincompatibility and myophily in Octomeria (Orchidaceae, Pleurothallidinae) species. Pl. Syst. Evol. 283: 1-8. Bawa, K. S. 1974. Breeding systems of tree species of a lowland tropical community. Evolution 28: 85-92. Bawa, K. S. 1979. Breeding systems of trees in a wet forest. New Zealand J. Bot. 17: 521-524. Borba, E. L. 2003. Novas combinaes em Acianthera (Pleurothallis s.l.; Orchidaceae: Pleurothallidinae) ocorrentes nos campos rupestres brasileiros. Sitientibus, Sr. Cincias Biol 3: 22-25. in three Bulbophyllum (Orchidaceae) species occurring in the Brazilian campos rupestres. Lindleyana 13: 203-218. Borba, E. L. & J. Semir. 1999. Temporal variation in pollinarium size after its removal in species of Bulbophyllum : a different mechanism preventing selfpollination in Orchidaceae Pl. Syst. Evol. 217: 197-204.
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Phylogenetic relationships in Pleurothallidinae (Orchidaceae): combined evidence from nuclear and plastid DNA sequences. Amer. J. Bot. 88: 2286-2308. Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 2005. Genera Orchidaceaearum. Volume 4. Epidendroideae (Part one). Oxford University Press, Oxford, UK. Proctor, M., P. Yeo & A. Lack. 1996. The natural history of pollination Harper Collins, London, UK. Richards, A. J. 1997. Plant breeding systems. 2nd ed. Chapman & Hall, London, UK. Scacchi, R. & G. De Angelis. 1989. Isoenzyme polymorphisms in Gymnadenia conopsea and its inferences for systematics within this species. Biochem. Syst. Ecol. 17: 25-33. Scacchi, R., G. De Angelis & P. Lanzara. 1990. Allozyme variation among and within eleven Orchis species (fam. Orchidaceae), with special reference to hybridization aptitude. Genetica 81: 143-150. Schlegel, M., G. Steinbrck, K. Hahn & B. as revealed by enzyme electrophoresis. Pl. Syst. Evol. 163: 107-119. Singer, R. B. & A. A. Cocucci. 1999. Pollination mechanism in four sympatric southern Brazilian Epidendroideae orchids. Lindleyana 14: 47-56. Sobrevilla, C. & M. T. K. Arroyo. 1982. Breeding systems in a montane tropical cloud forest in Venezuela. Pl. Syst. Evol. 140: 19-37. Tan, K. H., R. Nishida & Y. C. Toong. 2002. Floral synomone of a wild orchid, Bulbophyllum cheiri lures Bactrocera 28: 1161-1172. Tan, K. H., L. T. Tan & R. Nishida. 2006. Floral phenylpropanoid cocktail and architecture of Bulbophyllum vinaceum Chem. Ecol. 32: 2429-2441. Tremblay, R. L., J. D. Ackerman, J. K. Zimmerman & R. N. Calvo. 2005. Variation in sexual reproduction in orchids 84:1-54. van den Berg, C., D. H. Goldman, J. V. Freudenstein, A. M. Pridgeon, K. M. Cameron & M. W. Chase. 2005. An overview of the phylogenetic relationships within Epidendroideae inferred from multiple DNA regions and recircumscription of Epidendreae and Arethuseae (Orchidaceae). Amer. J. Bot. 92: 613-624. van den Berg, C., E. E. Higgins, R. L. Dressler, W. M. Whitten, M. A. Soto-Arenas & M. W. Chase. 2009. A phylogenetic study of Laeliinae (Orchidaceae) based on combined nuclear and plastid DNA sequences. Ann. Bot. 104: 417-430. pollination and evolution University of Miami Press, Coral Gables, Florida, USA. Zapata, T. R. & M. T. K. Arroyo. 1978. Plant reproductive ecology of a secondary deciduous tropical forest in Venezuela. Biotropica 10: 221-230.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. BORBA et al. Mating systems in the Pleurothallidinae221
on Andean Orchids was held in Ecuador in 2009. This country of only 256,371 square kilometers (98,985 the east across the Andes mountains and ultimately to the Galapagos Islands 972 km (604 miles) west of the Ecuadorian mainland. Indeed, because of their high LANKESTERIANA 11(3): 223. 2011.A LOOK AT THE ORCHID BOOK IN CELEBRATION OF CHARLES DARWINS 200TH BIRTHDAY KENNETH M. CAMERON Madison, Wisconsin 53706 U.S.A. email@example.com ABSTR A CT. the Origin of Species On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects their pollinators. The nine chapters treated members of Orchideae, Arethuseae, Neottieae, Vanilleae, Malaxideae, Epidendreae, Vandeae, Cymbidieae (especially Catasetum and on the Good Effects of Intercrossing RESU M EN. orqudeas, ya que fue el primer libro publicado despus del Origen de las Especies Sobre las estrategias por las cuales las orqudeas britnicas y las introducidas son fertilizadas por insectos Vandeae, Cymbidieae (especialmente Catasetum y sobre el Buen Efecto del Entrecruzamiento KEY WORDS :
1831-1836, the young English naturalist Charles HMS Beagle both the eastern and western coastlines of South America. It was during his time among the islands detailed and published in 1859 exactly 150 years birthday (he was born February 12, 1809). For these biology in particular, are worthy of consideration this to do so in Ecuador. The young naturalist/botanist well known, but the public is generally ignorant of further by regular interactions with his teachers, friends, and colleagues who shared a similar passion Professor of Mineralogy after becoming Professor of Botany, a subject he held in higher regard and him into the study of natural history and also for introducing him to such wellrespected botanists as Botanic Gardens, Kew, for 20 years. At Kew, Charles to the greatest American botanist of the 19th century, professional relationship through regular written correspondence. There is no doubt that the shared passion for botany among all of these men cemented pursuits. natural selection is to be found in his Journal and Within the pages of this best-selling book it is well known that he documented the fact that different bird species inhabited different islands in the archipelago, with some of the endemic plant species that he encountered: the aboriginal plants of the different islands wonderfully different... Scalesia, a remarkable to the archipelago: it has six species; one from Chatham, one from Albemarle, one from Charles Island, two from James Island, and the sixth from one of the three latter islands: not one of these six species grows on any two islands. The species of the Composit are particularly local; in like of tortoise, and of the widely distributed American genus of the mocking-thrush, as well as of two of make his greatest contribution to science by publishing On The Origin of Species important, and the reader is encouraged to explore the its importance to human society. After being thrust into did not end his basic research. In fact, he continued to publish until his death in 1882. It is generally underLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.224 LANKESTERIANA
and orchids (see Table 1). In fact, the next book to be published immediately after the Origin was his 300page treatise, On the Various Contrivances by which LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CAMERON 225 TABLE boldface: 9 of 21=43%.1839: Journal and Remarks (The Voyage of the Beagle) 1842: The Structure and Distribution of Coral Reefs 1844: Geological Observations of Volcanic Islands 1846: Geological Observations on South America 1849: Geology from Prepared for the Use of Her Majestys Navy: and Adapted for Travellers in General., John F. W. 1851: A Monograph of the Sub-class Cirripedia, with Figures of All the Species. The Lepadidae; or, Pedunculated Cirripedes. 1851: A Monograph on the Fossil Lepadidae; or, Pedunculated Cirripedes of Great Britain 1854: A Monograph of the Sub-class Cirripedia, with Figures of All the Species. The Balanidae (or Sessile Cirripedes); the Verrucidae, etc. 1854: A Monograph on the Fossil Balanid and Verrucid of Great Britain 1858: On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection (Extract from an Unpublished Work on Species) 1859: On the Origin of Species by Means of Natural Selection 1862: On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects 1868: Variation of Plants and Animals Under Domestication 1871: The Descent of Man, and Selection in Relation to 1872: 1875: Movement and Habits of Climbing Plants 1875: Insectivorous Plants 1876: The Effects of Cross and Self Fertilisation in the Vegetable Kingdom 1877: The Different Forms of Flowers on Plants of the Same Species 1880: The Power of Movement in Plants 1881: The Formation of Vegetable Mould Through the Action of Worms British and Foreign Orchids are Fertilised by Insects among the pages of this text, specialists in other and on the Good Effects of Intercrossing Inbreeding and outcrossing would later write an entire book on the subject Effects of Cross and Self Fertilisation in the Vegetable Kingdom (Moore, 2005) that he had a personal interest in the 62 aunts, uncles, and cousins born in the four generations and Sarah Wedgwood, 38 (61%) remained childless. did not reproduce. Perhaps some of these couples chose to lack of modern medical care (although the families children produced grandchildren for Charles and Emma. precisely the case (Moore, 2005).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.226 LANKESTERIANA publishing his orchid book to demonstrate that natural plants tend to self-pollinate (inbreed) more than do animals, because the former are stationary and their argued that outcrossing among plants must be more demonstrate that these, and most likely other plants fact, when you think about it, what better group of organisms to document this fact than orchids, which organs are intimately united into one. This is certainly to follow in most cases from cross-fertilisation, it is demonstrates that there must be something injurious tells us, in the most emphatic manner, that she abhors runs through the entire orchid book, the fact that Origin of Species as well. In his rush to publish the Origin as quickly as material that he felt bolstered his argument. Critics but also for so-called natural theologians who were beginning to accept many of the facts presented by geologists, paleontologists, and biologists in terms of the origins and antiquity of life but still saw God as the force guiding all laws of nature. Just as people either by means of natural selection or the hand of a creator, so they did as well in the mid-19th century as the Origin became a bestseller. Throughout the Spiranthes autumnalis before Among the most celebrated of the detailed examples hypothesis concerning the pollination of Angraecum sesquipedale. This orchid species endemic to epithet of the species, sesquipedale, translates to pollinator of the comet orchid (Xanthopan morganii praedicta astonishing length of the nectary had been acquired Madagascar became larger through natural selection in relation to their general conditions of life [or their plants of the Angraecum which had the longest some Orchids) ... would be best fertilised. These plants yield the most seed, and the seedlings would generally inherit long nectaries; and so it would be in
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CAMERON 227The orchid book, chapter by chapter read for any botanist, and especially for those of us interested in orchidology. Yam et al. (2009) examined other than pollination ecology, such as physiology, detailed account of how the book came to be, and the reader who desires further information is encouraged of orchids, and included 34 illustrations dispersed taxa, especially from the Neotropics, Australia, and temperate North America. These included Pterostylis, Caladenia, Thelymitra, Disa, Gongora, Sobralia, Pogonia, Platanthera, and others. In total, the second edition was expanded to treat 85 different genera of orchids (a 35% increase), four additional illustrations into nine. For the most part, these were arranged day as proposed by John Lindley (1827). Subfamilies were not considered, but Orchidaceae (excluding Chapters 1 and 2 treat Ophreae, in particular the European terrestrial orchids such as Orchis and Ophrys, the bee orchids. A total of 73 pages were most familiar because they grew near his home in the English countryside. Among the genera considered are Orchis, Ophrys, Herminium, Peristylus, Gymnadenia, Platanthera, Habenaria, Disa, and Bonatea. Chapter 3 and 4 are short and consider tribes Vanilla these lower epidendroid groups, it was treated among the 12 pages of Chapter 3. Among the other genera discussed here are Cephalanthera, Sobralia, Pterostylis, Caladenia, and Pogonia. More recent systems of (e.g., Chase et al. orchids, which share plesiomorphic aspects of their subfamilies. Likewise, many of the genera considered Epipactis, Listera, Neottia, Epipogium, Goodyera, Spiranthes, Thelymitra of subfamilies Orchidoideae and Epidendroideae. The subsequent chapters of the orchid book treated largest subfamily of Orchidaceae. Today we count Lindley estimated that there might be as many of 6000 and Epidendreae. These are primarily tropical orchid tribes, and many of the most species-rich genera were still poorly collected, especially those from higher FIGURE The Various Contrivances by which Orchids are Fertilised by Insects. 2nd edition.John Murray, London.
primarily through relationships with horticulturists at Kew and from other glasshouse collections were Laelia, Epidendrum, Pleurothallis, and Stelis Zootrophion atropurpureum which he knew and illustrated in Figure 20 as Masdevallia fenestrata. The taxonomy below as an Appendix. The names as they appeared in the second edition of the orchid book as well as their Following in sequence we reach Chapters 6 and (as then understood), and particular emphasis was Catasetum and its fast-action mechanism of pollinarium ejection. This is not surprising because In 1875, he published Movement and Habits of Climbing Plants Drosera Dionaea muscipula Ellis). In 1880, he published The Power of Movement in Plants. Other orchids discussed in chapters 6 and 7 of the orchid book are Calanthe, Miltonia, Sarcanthus, and Cycnoches. with one genus (Cypripedium into tribe Cypripedieae. Among the 20 pages is also found a discussion concerning the homologies of and experimental biologist. For example, he described Cypripedium calceolus in order to record proboscis, as I had supposed, from the outside through contrary to the argument made throughout the book in so he was delighted to document that his experiments Cypripedium also manages An example of a hypothesis presented in the book can be found in this chapter. Figure 36 in the book anthers of an ancestral inner androecial whorl were the column. One of the anthers from the outer whorl whorl combined with the lower petal, forming the its construction. naturally be inquired, Why do the Orchideae exhibit so sure that many other plants offer analogous adaptation of high perfection; but it seems that they are really more numerous and perfect with the Orchideae than with most who was passionate about animal and plant biology and us today feel the same and should feel pride, but also humility, in following his footsteps.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.228 LANKESTERIANA
APPENDIX and spellings are reproduced exactly as they appear within the index of the book. Names within brackets are those accepted currently by the World Checklist of Monocotyledons. (2009). The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the Internet; http://www.kew.org/wcsp/monocots/ accessed 2 August 2009.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CAMERON 229Aceras anthropophora, 26, 258 [Orchis anthropophora] longibracteata, 26 [Barlia robertiana] 90 fornicatus, 90, 280 sinclairii, 90, 280 Acontia luctuosa, 31 [Acronia luctuosa = Pleurothallis luctuosa] Acropera, 154, 156, 276 [Gongora] loddigesii, 166 [Gongora galeata] luteola, 166, 239 [Gongora galeata] Aerides, 156, 265 cornutum, 265 [Aerides odorata] odorata, 158 virens, 156 [Aerides odorata] Angrcum, 251 distichum, 154 eburneum, 155 sesquipedale, 154, 162, 282, 265 Apostasia, 248 146 Bolbophyllum, 274, 276 [Bulbophyllum] barbigerum, 138 cocoinum, 137 cupreum, 137, 265 rhizophor, 137 [Bulbophyllum falcatum velutinum] Bonatea speciosa, 71, 76, 244, 264, 361 Brassia, 156 Caladenia dimorpha, 89 Calna, 89 [Caleana] Calanthe dominii, 161 [Calanthe x dominii] masuca, 161, 267, 269 [Calanthe sylvatica] veratrifolia, 280 [Calanthe triplicata] vestita, 162 Catasetum, 256, 270 callosum, 192, 195 luridum, 191 mentosum, 206 planiceps, 193 saccatum, 180, 239 tabulare, 192 tridentatum, 191, 196, 197, 239, 256, 269 [Catasetum macrocarpum] Cattleya, 143 239, 265 crispa, 147 [Sophronitis crispa] Cephalanthera, 277 ensifolia, 86 [Cephalanthera longifolia] 80 239, 242, 249, 259, 269, 277, 287, 290 [Cephalanthera longifolia] Chysis, 146 Cirrha, 171 Coelogyne cristata, 146 Coryanthes, 90, 173, 232, 265 175 macrantha, 175 speciosa, 174 triloba, 281 [?] Cycnoches egertonianum, 224 ventricosum, 220 Cymbidium giganteum, 155, 252, 260, 263 [Cymbidium iridiodes] Cypripedium, 226, 229, 262, 275 acaule, 229 barbatum, 239 [Paphiopedilum barbatum] calceolus, 229, 282 candidum, 235 pubescens, 229, 230 [ pubescens] purpuratum, 239 [Paphiopedilum purpuratum] Cyrtostylis, 90 Dendrobium, 287 bigibbum, 142 chrysanthum, 138, 265 cretaceum, 142, 291 [Dendrobium polyanthum] formosum, 142 speciosum, 281 tortile, 142 Disa, 265 cornuta, 78 77, 281  macrantha, 78, 290 [Disa cornuta] Disperis, 265
Epidendrum cochleatum, 249 [Prosthechea cochleata] 146, 249 [Epidendrum paniculatum] glaucum, 146 [Dichaea glauca] Epipactis, 239, 251 latifolia, 100, 101, 259, 282, 287 [Epipactis helleborine] microphylla, 102 palustris, 93 275 purpurata, 102 rubiginosa, 102 [Epipactis atrorubens] 102, 291 [Epipactis purpurata] Epipogium gmelini, 103 [Epipogium aphyllum] Eulophia viridis, 156, 269 [? = Eulophia epidendraea] Evelyna, 265 [Elleanthus] carivata, 146, 239, 241 [Elleanthus caravata] 155 [Galeandra baueri] Glossodia, 237 Gongora, 276 atropurpurea, 169 maculata, 168 truncata, 169 Goodyera, 239, 260 discolor, 105 [Ludisia discolor] Goodyera pubescens, 105 repens, 103, 105 Gymnadenia, 251 albida, 43, 68 [Pseudorchis albida] conopsea, 32 40, 43, 65, 238, 239, 255, 271, 272 odoratissima, 68 tridentata, 68, 291 [Platanthera clavellata] Habenaria bifolia, 78, 40, 43, 251 [Platanthera bifolia] Habenaria chlorantha, 43, 69, 239, 244, 251 [ ] Herminium monorchis, 59, 61, 255 Llia, 146 cinnabarina, 148 [Sophronitis cinnabarina] Leptotes, 146 Liparis pendula, 239, 241  Listera, 251, 287 [Neottia] cordata, 124 [Neottia cordata] ovata, 115, 276 [Neottia ovata] 155, 260 251, 276 paludosa, 32, 129, 239, 241, 241, 258, 284 [ Hammarbya paludosa] Masdevallia, 241, 274, 276 fenestrata, 135, 136, 142 [Zootrophion fenestratus] 156, 278 ornithorhyncha, 157, 159 [?] Megaclinium falcatum, 138 [Bulbophyllum falcatum] Microstylus rhedii, 132, 135  Miltonia clowesii, 154, 155 Monachanthus viridis, 196, 197, 198, 201 [Catasetum cernuum] Mormodes ignea, 208-219, 249, 276, 283 219 Myanthus barbatus, 192, 199, 203, 205 [Catasetum barbatum] Neotinia intacta, 27, 291 [Neotinea maculata] Neottia nidus-avis, 125, 258, 290 Nigritella angustifolia, 27 [Gymnadenia nigra] Notylia, 171 Odontoglossum, 156 Oncidium, 153, 156, 158, 239, 251, 266 unguiculatum, 252 Ophrys apifera, 52, 54-58, 259, 279, 291 arachnites, 51 [Ophrys apifera] aranifera, 50, 280 [Ophrys sphegodes] muscifera, 32, 45, 49, 280 [Ophrys insectifera] 52, 292 Orchis fusca, 15, 35, 37 [Orchis purpurea] hircina, 25, 39, 273 [Himantoglossum hircinum] latifolia, 15, 35, 37, 255 [Dactylorhiza incarnata] maculata, 15, 34, 32 35, 37, 39, 255, 255, 277, 278 [ Dactylorhiza maculata] Orchis mascula, 6, 273, 278 militaris, 36, 37 morio, 15, 128, 33, 37, 39, 278 [Anacamptis morio] pyramidalis, 16, 21, 34, 37, 39, 38, 254, 256, 260, 261, 264, 272, 273 [Anacamptis pyramidalis] ustulata, 25 [Neotinea ustulata] Ornithocephalus, 160 Peristylus viridis, 43, 63, 255 [Coeloglossum viride] Phaius, 146 grandifolius, 280  Phalaenopsis, 153, 159, 276 amabilis, 159 159, 269 [Phalaenopsis amabilis] Platanthera, 75 chlorantha, 69 dilatata, 77 [Piperia dilatata] 76, 77 75LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.230 LANKESTERIANA
hyperborea, 76, 291 Pleurothallis ligulata, 135 [Stelis ligulata] prolifera, 135 [Acianthera prolifera] Pogonia ophioglossoides, 86 Pterostylis, 232 87, 89 [? Pterostylis longifolia] trullifolia, 86, 88, 280 Rodriguezia secunda, 159 [Rodrigueza lanceolata] suaveolens, 156, 159 [Gomesa foliosa] Saccolabium, 153, 156 Sarcanthus, 276 [Cleisostoma] parishii, 142 [Cleisostoma parishii] teretifolius, 154, 156, 268 [Cleisostoma simondsii] Selenipedium palmifolium, 232 Serapias cordigera, 27 Sobralia macrantha, 91 Sophronitis, 146 Spiranthes australis, 114, 275, 291 [Spiranthes sinensis] autumnalis, 106-114, 239 cernua, 111 gracilis, 111 [ Chlorosa gracilis] Stanhopea, 155, 276 devoniensis, 171 [Stanhopea hernandezii] oculata, 171 Stelis, 274 135  Thelymitra, 291 carnea, 127, 280 127 [? Thelymitra longifolia] Uropedium, 240 [Phragmipedium] Vanilla aromatica, 90 [Vanilla planifolia] Warrea, 155, 270 155 [Zygopetalum maculatum]LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CAMERON 231 LITERA TURE CITED in Orchid conservation. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life John Murray, London, UK. in M. Porter, and M. Richmond (eds.). The correspondence of Charles Darwin in: F. Burkhardt, J. Browne, The correspondence of Charles Darwin in: F. Burkhardt, J. Browne, The correspondence of Charles Darwin in: F. Burkhardt, J. The correspondence of Charles Darwin Press, UK. On the various contrivances by which British and foreign orchids are fertilised by insects and the good effects of intercrossing. John Murray, London, UK. On the various contrivances by which British and foreign orchids are fertilised by insects, 2nd ed. John Murray, London, UK.Lindley, J. 1830. The genera and species of orchidaceous plants. Ridgways, London, UK. World Checklist of Monocotyledons. (2009). The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the Internet; http://www.kew.org/wcsp/monocots/ accessed 2 August 2009.
Almost half a century ago, the late William Stearn (1960), addressing the 3rd World Orchid Conference in London, presented what he considered to be 10 landmarks in the knowledge of orchids, a family which he labeled as the most promiscuous of all plants based on the ability of often unrelated species to produce Romans dating from before the time of Christ. in late 17th century. orchids to Europe in the late 17th and early 18th centuries. 4. The application of binomial nomenclature to orchids by Linnaeus in 1753. 5. The change in method for heating greenhouses from dry to wet heat, stimulating their introduction about them in the 19th century. 6. The elucidation of the pollination mechanisms of hybrid in 1861.LANKESTERIANA 11(3): 233. 2011.ORCHIDS IN A CHANGING CLIMATE PHILLIP CRIBB Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom ABSTRACT. two millennia or more, our knowledge of orchids remained sketchy, mainly because the main centers of learning rapidly. Almost half a century ago, Professor William Stearn outlined this progress in his landmark lecture at RESU M EN. hito, presentada en la Tercera Conferencia Mundial de Orqudeas en Londres. Sin embargo, el conocimiento un efecto dramtico sobre las orqudeas del mundo. KEY WORDS :
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.234 LANKESTERIANA intergeneric hybrid in 1868. association. asymbiotically in the early 20th century. past few years that I could easily list many more. The new major landmarks I would submit are as follows: worldwide trade in orchid hybrids. checklist of orchids on the internet. orchid systematic problems, including estimating the age of the family. Publication of Genera Orchidacearum, introducing a new system of molecular and morphological data. allowing dating of the origins of the family. orchids and their habitats and the need for their in situ and a situation made more urgent by the threat of climate change. I will deal with each in turn, some in more detail than Micropropagation Cloning orchids by meristem culture is so commonplace nowadays that we tend to forget what transformed from the playthings of the rich to an charismatic of plants has been bolstered. Not core orchid community. Orchids are now commonly sold without their correct name. Plants are marketed without a grex or clonal name. The introduction of plant patents some years ago has further confused to those clones that differ from the parental plant. The problem does not, of course, affect the buyer whose produce new hybrids. If the parental names are absent or incorrect, the offspring cannot be named according (Brickell, 2004). It could be argued that names are unimportant, but, as I will elaborate upon later, they form the backbone of access to knowledge about orchids, indeed about all organisms. Computing changed the world. I would like to consider how they holistic approach, but the quality of information American Orchid Society (www.aos.org) and Royal high quality information and images on many aspects of orchids. A number of societies and orchid groups also publish their journals and newsletters on the web. Original high-quality information on orchids can also be sought on a number of other websites. I would World Monocot Checklist (http://www.kew.org/wcsp/ monocots) based at the Royal Botanic Gardens, Kew. It is regularly updated through an international network
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CRIBB Orchids in a changing climate235 Why is a checklist of orchids important? We cannot communicate satisfactorily about anything unless the only reliable spine to access information on orchids. The second website I would recommend and use unibas.ch/site.herbarium. php). Features of this website include thousands of images of orchids (photographs, illustrations from all of the historically important orchid books, and herbarium specimens) and access to the most complete bibliography of the orchids (BibliOrchidea). A searchable biographic database another useful feature. DNA analysis It would not be an exaggeration to state that the ability our understanding of the family. The work of Mark Chase and his many collaborators has resulted in the marked contrast to the position just 25 years ago when few scientists were attracted to the family, which was considered to be too large and too horticultural to be of interest. In short, most scientists preferred to work on smaller families where their results were less likely to This situation has fortunately changed dramatically. collaboration of some of the brightest young scientists. Their work has led to a better understanding of orchids as a family, the relations of its constituent parts, and We now know that: asparagoid monocots (Table 1), of which the best known is a terrestrial genus with plicate 2) Orchids are a monophyletic family that includes apostasioids, cypripedioids, and the rest of considered discrete families by some authors (e.g. Rasmussen, 1985). 3) Vanilla ancient lineage worthy of subfamilial status. 4) Spiranthoid orchids are not worthy of subfamilial status and comprise a group within the orchidoids. 5) The circumscriptions of many long-accepted genera, e.g. Cattleya, Laelia, Masdevallia, and Oncidium notably Odontoglossum, do not warrant recognition at all. These ideas are currently being assembled in the monumental Genera Orchidacearum (Pridgeon et al. TABLE 1. Families of Asparagales allied to Orchidaceae (Chase, 2001).
Warner, which is seeking to barcode the 1300 or so orchid species found in the country. The consequences of this for an understanding of species delimitation, to be considered. populations of the Western Australian underground orchid (Rhizanthella gardneri) might represent two Australian orchid genera, notably and Brown, 2007) and Chiloglottis (Florian Schiestl, personal communication). accepted European Ophrys probable that many new species are described where the changing of generic concepts from long-accepted ones. This affects a number of the most important genera in horticulture, including Cattleya, Laelia, Masdevallia, Odontoglossum, and Oncidium. In the decades in horticultural use when the botanists had long since consigned them to synonymy. For the past few years or so, the system has been abandoned, been used by the registrar, leading to many changes (nothogeneric) names. The rationale for this is that to different interpretation, so there is plenty of room for disagreement. My own opinion is that the present system of nothogeneric recognition is no longer useful which use a personal surname followed by -ara. I system will simplify registration and label writing as long as grex names are not repeated within a breeding group (which they mostly are not!). Mark Chase, Sarah Thomas, and I spelled out the need for a new system some years ago (Cribb et al., 1999). The orchid fossil record Meliorchis caribea et al. (2007) in the journal Nature. It comprised an orchid pollinarium on the back of a bee, the extinct Proplebeia dominicana, set in 15-20 million-year-old Miocene our knowledge of the antiquity of the orchids. The pollinarium can be safely assigned to a species of the terrestrial Goodyerinae, possibly Kreodanthus or Microchilus of particular orchid taxa can be estimated from the sequences. Chase (2001) suggested that the orchid lineage might be up to 90 million years old, in contrast et al. (2007) suggested a date of about 76-84 million years ago in the late Cretaceous for the emergence of the family. Both support a pattern of an ancient family contrast, the predominantly terrestrial orchidoids and LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.236 LANKESTERIANA
recent times, particularly since the end of the last glaciation in the tropical mountain chains, such as the and the mountains of the Malay Archipelago. Conservation The orchid world can be proud of its considerable last 20 years. It has been aware of the rarity of many species, especially some of the showiest orchids, expressed concern at the scale of orchid collection for increasing rate at which orchids and their habitats to address these serious issues. The causes are well appreciated by the public at large. The Orchid Specialist Group, an arm of IUCN successful in stimulating research and projects on endangered orchids. The OSG comprises some 200 orchid scientists and horticulturists worldwide. Under the chairmanship of Michael Fay of the Royal Botanic Gardens, Kew, it has sponsored three successful Australia, USA, and Costa Rica. The fourth is due in information can be gained from the OSG website also produces an electronic newsletter. Perhaps the main result of the work of the OSG has been to bring together current ideas and methodologies the main products of the First Conference in Perth, Australia, was a techniques manual entitled Orchid et al., 2003). In 2003, the OSG established a charitable foundation the American Orchid Society and Australian Orchid On a broader scale, I would like to mention the recent in Chengdu, China, and Quito, Ecuador, attracted a broad-based response. The project, Orchid Seed Stores for Sustainable Use, aims to establish protocols for orchid seed collection and storage based upon sound orchid seed-banks in orchid-rich countries. [See paper Orchids face increasing threats to their existence, population, logging, mining, and exploitation. Climate Roberts (personal communication) has shown that decrease and disappearance of orchids from local habitats that appear to be still suitable. Was this and bacteria that control germination and early growth IPCC, 2007). Conclusions years has meant that our knowledge of orchids has increased at a rate far greater than at any time in the been welcome in the orchid community, particularly those that require the relearning of plant relationships, being ditched and new ones proposed with frightening we can take some new concepts happily on board, LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. CRIBB Orchids in a changing climate237
secure future for orchids if is it is applied more widely. orchids, but often the limiting factor is funding. The orchids is one that is good news for orchids and orchid be found in the wild or are protected and cannot be unknown effect of global climate change predicted by the recent report from a UN panel of experts. Will to hear the update on orchid landmarks in 50 years. LITERA TURE CITED Chase, M. W. 2001. The origin and biogeography of Orchidaceae. Pp. 1-5 in: A.M. Pridgeon, P. J. Cribb, M. W. Chase & F. N. Rasmussen (eds.) Genera orchidacearum, Volume 2. Orchidoideae (Part one) Press, UK. Cribb, P. J., Chase, M. W., and Thomas, S. 1999. Orchid hybrid generic names time for a change? Pp. 395-396 in: S. Andrews, A. Leslie, and C. Alexander (eds.). of cultivated plants Royal Botanic Gardens, Kew, UK. 2003. Orchid conservation Kota Kinabalu, Sabah, Malaysia. 2007. http://www.ipcc.ch/ hammer orchids (: Orchidaceae), with some pollinators. Austral. Syst. Bot. 20: 252-285. Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 1999. Genera orchidacearum, Volume 1. General introduction, Apostasioideae, Cypripedioideae Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 2001. Genera orchidacearum, Volume 2. Orchidoideae (Part one) Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 2003. Genera orchidacearum, Volume 3. Orchidoideae (Part two), Vanilloideae Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 2005. Genera orchidacearum, Volume 4. Epidendroideae (Part one) Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen. 2009. Genera orchidacearum, Volume 5. Epidendroideae (Part two) from a fossil orchid with its pollinator. Nature 448: 10421045. Rasmussen, F. N. 1985. Orchidaceae. Pp. 249-274 in: R. The families of monocotyledons: structure, evolution and Springer Verlag, Berlin, Germany. Schmid, R. and Schmid, M. J. 1977. Pp. 17 in: J. Arditti (ed.). Orchid biology: reviews and perspectives 1. Cornell Stearn, W. T. 1960. Two thousand years of orchidology. Pp. 26-41 in: P. M. Synge (ed.). Proceedings of the 3rd World Orchid Conference UK. Willis, J. C. 1973. ferns238 LANKESTERIANA
especies de Sobralia. Ahora tenemos algunas otras en preparacin, y media docena que se pueden publicar tan quiero discutir algunas poblaciones a las cuales Sobralia son sumamente uno puede encontrar plantas que concuerden bien con las descripciones originales. Las primeras especies de Sobralia de Costa Rica y Panam fueron descritas por Reichenbach en 1852. La mayora de ellas son ntidas y fciles de reconocer, con la excepcin notoria de S. bletiae Rchb.f. El tipo de S. bletiae tiene un ptalo ligeramente adherido al labelo en la base (afortunadamente, este holotipo se un lbulo lateral, y en su dibujo agreg otro igual al describi como S. suaveolens Rchb.f. Sobralia warszewiczii tambin fue descrita en 1852, y el nombre se ha usado desde Panam hasta una muestra prensada ms bien fea, probablemente concuerdan muy bien con el ejemplar prensado. En esa a aproximadamente 2000 m de altura en el Volcn Sobralia warszewiczii plantas de S. warszewiczii en Costa Rica, pero es muy est muy cerca de la frontera tica. LANKESTERIANA 11(3): 239. 2011. SOBRALIA WARSZEWICZII ROBERT L. DRESSLER firstname.lastname@example.org RESU M EN. Sobralia warszewiczii Rchb.f. fue descrita de Panam occidental en 1852. El nombre se ha usado hasta del todo con S. warszewiczii. Una especie que forma colonias grandes en el noroeste de Costa Rica podra ser la misma especie que se encuentra en el sur de Mxico, pero requiere ms estudio, y no son nada claros los S. labiata S. warszewiczii). ABSTRACT. Sobralia warszewiczii Rchb.f. Rchb.f. was described from western Panama in 1852. The name has Costa Rica may well be the same species that occurs in southern Mexico, but more study is needed, and it is not clear what the correct names are for any of this complex (except for S. labiata S. warszewiczii swarms. PALABRAS CLA VE / KEY WORDS : Sobralia. S. warszewiczii, enjambres hbridos, hybrid swarms
Aparte de S. labiata que es muy distinta, dos supuestas especies se han descrito. Sobralia amparoae Schltr. fue pero no se sabe nada con respecto al origen de la planta. En el mismo artculo, Schlechter public Sobralia bradeorum Sobralia iguales a las que hay de encontrar Cypripedium S. bradeorum fueron daadas hasta Guatemala y el sur de Mxico, pero parece que Rica y Panam. S. amparoae tena cinco quillas, mientras que S. bradeorum tena solamente tres. Con respecto a las quillas, el complejo de S. amparoae, S. bradeorum y S. warszewiczii, por lo general, tienen 3, 4 o 5 quillas principales, ms una quilla ms baja a cada lado. Me parece que hay son las poblaciones que podran representar enjambres hbridos. Otra caracterstica a la que se ha dado mucha importancia es que plantas que crecen a mayor altura o en lugares muy expuestos tienen las hojas ms bien coriceas y fuertemente acanaladas, pero si se caractersticas desaparecen. grandes de una Sobralia en estructura. Las poblaciones grandes de Guanacaste es muy limitada. Al parecer, las poblaciones grandes y Chiapas bien podran ser la misma especie que forma (adems de S. labiata extiende desde Coto Brus hasta el Valle de Oros, en Cartago. Suponiendo que la planta de doa Amparo fue trada por su hermano de Turrialba, es muy posible que S. amparoae sera el nombre correcto para esta especie (Fig. 4). 2. Por la carretera al sur de Cartago, cerca de Cangreja (km. 3035) hay una Sobralia de otras especies muy diferentes. a apenas unos 10 km al sureste del Jardn Botnico Lankester, pero hay que bajar al Valle de Oros y subir pero llena de sobralias creciendo en los rboles y plantas de S. amparoae y otras muy oscuras con el creciendo muy bien (ya tenemos plntulas en frasco.) Sera muy interesante hacer un estudio detallado de la poblacin del Alto de Araya, pero la condicin de la carretera no ayuda mucho. Tambin hay lo que parece ser otro enjambre hbrido en La Laja, ms al sur, tambin en Orosi. Afortunadamente, La Laja es de ms fcil acceso, y creo que ser mucho ms factible hacer un estudio detallado con las plantas de La Laja. frecuentemente forman colonias grandes a los lados S. ecuadorana S. gentryi S. powellii LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.240 LANKESTERIANA
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. DRESSLER El complejo de Sobralia warszewiczii241 FIGURE 1. Sobralia warszewiczii, una planta recolectada en el area de Ro Sereno, Panam, cerca de la frontera con Costa FIGURE 2. Sobralia sp., Esta especie se encuentra cerca de Cangreja, al sur de Cartago, Costa Rica.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.242 LANKESTERIANA FIGURE Sobralia amparoae? Esta especie se encuentra de Coto Brus (Puntarenas, Costa Rica) hasta el rea de Tapant cuales otras especies contribuyan genes a los enjambres hbridos. FIGURE 3. Sobralia
Schltr. forman un enjambre hbrido a los lados de las carreteras, aunque las mismas especies no parecen formar hbridos en regiones no alteradas. Es muy S. powellii S. powellii LITERA TURA CIT ADA Ecuador. Fascculo 6. Orquideologa 21: 3-67. 715. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. DRESSLER El complejo de Sobralia warszewiczii243 FIGURE
Scaphosepalum agrupa a 49 especies y es un clado subtribu Neotropical Pleurothallidinae la que cuenta con aproximadamente 4000 especies (Pridgeon et al., 2005). Scaphosepalum puede ser fcilmente coronados por osmforos (Fig. 1), los que en la mayora de las especies, estn bien desarrollados (Vogel, 1965; Pridgeon & Stern, 1985; Luer, 1986, 1988; Pridgeon et al., 2005). Los sinspalos y el spalo dorsal poseen inicialmente se incluy a este gnero en Masdevallia (Luer, 1986). Scaphosepalum y de la subtribu Pleurothallidinae se encuentra en los bosques montanos del norte de los Andes, un paisaje Graham, 1999; Young et al., 2006; Rull, 2008; Antonelli et al., 2009; Graham, 2009; Struwe et al., Scaphosepalum para examinar los patrones de especiacin del gnero con el LANKESTERIANA 11(3): 245. 2011.FILOGENIA MOLECULAR PRELIMINAR DE SCAPHOSEPALUM (ORCHIDACEAE:PLEUROTHALLIDINAE) LORENA ENDARA A. 1,2,3 NORRIS H. WILLIAMS 1,2 Y W. MARK WHITTEN 11 Florida 32611-7800, U.S.A. 23 RESUMEN. El gnero Scaphosepalum (Orchidaceae: Pleurothallidinae) agrupa a 49 especies de distribucin tanto especies es la consecuencia de especiacin simptrica o especiacin aloptrica seguida por contacto secundario. Scaphosepalum trnL-F, matK, y ycf1. Los datos ABSTR A CT. The orchid genus Scaphosepalum (Orchidaceae: Pleurothallidinae) encompasses 49 species widely or narrowly distributed in biotic sympatry in the montane Neotropical forests, where it reaches its peak of Scaphosepalum and use the resulting phylogenetic hypothesis to determine if the current sympatric distribution of the species is the result of sympatric or allopatric speciation followed by secondary contact. The preliminary phylogeny presented here is based on trnL-F, matK, and ycf1. Analyses were performed using Maximum Parsimony (MP), Maximum Likelihood indicate a strong geographical structure in the dataset. PALABRAS CLA VE / KEY WORDS : Orchidaceae, Scaphosepalum
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.246 LANKESTERIANA FIGURA Scaphosepalum. A. S. decorum; B. Flor tubular de S. odontochilum ; C. S. medinae. osm: osmforos, ds: spalo. FIGURA Scaphosepalum
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. ENDARA et al. Filogenia preliminar de Scaphosepalum247 puntos calientes (hot-spots) del planeta (Myers et al., 2000). Scaphosepalum representa un sistema ideal para detectar escenarios de especiacin aloptrica, paraptrica y simptrica pues una gran cantidad de sus Materiales y mtodos Muestreo especmenes que representan a 28 especies con los protocolos citados en Whitten et al. (2007). Todos los en el Jardn Botnico de Atlanta (ABJ) y cuentan con especmenes (FLAS). Secuenciacin de genes molecular constituye el mayor reto de este proyecto trnL-F, matK, y ycf1) y un gen molecular (ITS) han demostrado ser polimerasa Sigma Jumpstart Taq y Sigma Jumpstart redTaq (Sigma-Aldrich, St. Louis, Missouri, USA) en agua destilada. matK y ycf1 de los diferentes genes se detallan en las tablas 1 y 2, colocados en placas de 96 celdas y fueron secuenciados editados en el programa Sequencher 4.6 (Genecodes Inc., Ann Arbor, Michigan, USA) y alineados manualmente en el programa Se-Al (Rambaut, 2000). TABLA 1 Scaphosepalum.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.248 LANKESTERIANA TABLA 2 Scaphosepalum. Programas touchdown. (Farris et al., 1995) y pruebas de congruencia topolgica para cada set de datos (Soltis et al., 1998). El set de Mxima Parsimonia (MP), Mxima Verosimilitud (ML: Maximum Likelihood), y Bayesiano. Para los dos et al., 2004). Los Dryadella, Platystele, Teagueia, y Pleurothallis ( o Sarcinula) acanthodes sugeridos como grupos hermanos de Scaphosepalum (Pridgeon et al., 1999, 2005). El soporte para los nodos recuperados se generaciones y se corri el anlisis por seis millones de generaciones. Los anlisis fueron ejecutados en Florida. Resultados congruencia entre las regiones plstidas pero incongruencia entre los sets de datos plstidos y el set topologas de los rboles consenso estricto generados mediante MP y la topologa resultante de ML. La topologa del rbol correspondiente a las regiones
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. ENDARA et al. Filogenia preliminar de Scaphosepalum249 rbol nuclear. Estas diferencias fueron consideradas combinar los datos de las tres regiones plstidas y la regin nuclear. La Tabla 3 detalla las estadsticas de los rboles obtenidos en base a los anlisis parsimnicos combinadas (ITS, trnLF, matK, y ycf1) as tambin MP, ML, y anlisis Bayesianos son similares y Discusin Aunque en una etapa preliminar (28 especies con alto soporte bootstrap contiene prcticamente a todas las especies muestreadas de las estribaciones noroccidentales de los Andes y Amrica Central con potenciales casos de dispersin hacia las estribaciones orientales de los Andes centrales en Ecuador. Los dos clados restantes contienen a la mayora de las especies encontradas en las estribaciones oreintales de los Andes (color amarillo en l Figura 3). Es posible (Fig. 2), que las especies que habitan en los mismos hbitats no estn cercanamente relacionadas, en otras palabras, no son los grupos hermanos inmediatos, por ejemplo, Scaphosepalum beluosum, S. decorum, S. dodsonii, S. digitale, y S. ophiodon estn presentes en los mismos hbitats (Fig. 2) sin embargo no son TABLA 3 Scaphosepalum. Valores MP: Mxima Parsimonia.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.250 LANKESTERIANA FIGURA 3. Filogenia preliminar de Scaphosepalum matK, trnL-F, y ycf interno.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. ENDARA et al. Filogenia preliminar de Scaphosepalum251especies hermanas por lo tanto no son el resultado de especiacin simptrica. Scaphosepalum swertiifolium (Fig. 3), una especie 1998). A futuro se espera aumentar ms especmenes distribuidas y tambin especies que exhiben una distribucin (Scaphosepalum antenniferum, S. breve, S. microdactylum, y S. verrucosum) para explorar Llama la atencin la posicin de Scaphosepalum medinae como especie hermana al resto de Scaphosepalum. Scaphosepalum medinae (Fig. 1c) y S. pleurothallodes cimosas. Probablemente S. pleurothallodes tambin sea hermana al resto de Scaphosepalum morfolgica de S. pleurothallodes y S. medinae a Pleurothallis ( o Sarcinula) acanthodes, resulta interesante pues P. acanthodes crece en simpatra con las dos especies de Scaphosepalum antes mencionadas y podra ser el taxn hermano del clado de Scaphosepalum. Para comprobar la Scaphosepalum incluir a otras especies de Pleurothallis ( o Sarcinula). Es tambin una prioridad de nuestro proyecto un muestreo completo y denso de taxones ya a las estrategias de muestreo de taxones y caracteres paraptrica seguidos por contacto secundario de las especies. LITERA TURA CIT ADA Altekar, G., S. Dwarkadas, J. P. Huelsenbeck & F. Ronquist, 2004. Parallel Metropolis coupled Markov chain Monte Carlo for Bayesian phylogenetic inference. Bioinformatics 20: 407. Antonelli, A., J. A. A. Nylander, C. Persson & I. Sanmartin. 2009. Tracing the impact of the Andean uplift on Neotropical plant evolution. Proc. Nat. Ac. Sc. 106: 97499754. Burnham, R. J. & A. Graham. 1999. The history of Neotropical vegetation: new developments and status. Ann. Missouri Bot. Gard. 86: 546-589. Cladistics 10: 315319. phytogeographical connections between Central and accident of the Andean orogeny? Ann. Missouri Bot. Gard. 69: 557-593. Graham, A. 2009. The Andes: a geological overview from a biological perspective. Ann. Missouri Bot. Gard. 96: 371-385. Graybeal, A. 1998. Is it better to add taxa or characters to a Howard, D. J. & S. H. Berlocher (ed.). 1998. Endless forms: species and speciation Oxford University Press, UK. Huelsenbeck, J. P. & F. Ronquist. 2001. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754 755. Hughes, C. & R. Eastwood. 2006. Island radiation on a after uplift of the Andes. Proc. Nat. Ac. Sc. 103: 1033410339. Luer, C. 1986. Icones Pleurothallinidarum I Systematics of the Pleurothallidinae. Monogr. Syst. Bot. Missouri Bot. Gard. 15. Luer, C. 1988. Icones Pleurothallinidarum V Systematics of Dresslerella and Scaphosepalum. Monogr. Syst. Bot. Missouri Bot. Gard. 26. Myers, N., R. A. Mittmeier, G. A. B. da Fonseca & J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853-858. Posada, D. & K. A. Crandall. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817-818. Pridgeon, A. M. & W. L. Stern1985. Osmophores of Scaphosepalum (Orchidaceae). Bot. Gaz. 146: 115-123. Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen (eds.). 1999. Genera orchidacearum. Volume 1. General introduction, Apostasioideae, Cypripedioideae. Oxford University Press, UK. Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen (eds.). 2005. Genera orchidacearum. Volume 4. Epidendroideae (Part one). Oxford University Press, UK. Rambaut, A. 2002. Se-Al: Sequence Alignment Editor, ver. 2.0a11. URL: http://tree.bio.ed.ac.uk/ssoftware/seal.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.252 LANKESTERIANAReiseberg, L. H. & L. Brouillet. 1994. Are many plant species paraphyletic? Taxon 43: 21-30. Ronquist, F. & J. P. Huelsenbeck. 2003. MrBayes3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572. Rull, V. 2008. Speciation timing and neotropical biodiversity: the Tertiary-Quarternary debate in the light of molecular phylogenetic evidence. Molec. Ecol. 17: 2722-2729. Soltis, D. E., P. S. Soltis & J. J. Doyle. 1998. Molecular systematics of plants IIDNA sequencing Kluger Academic, Norwell, Massachusetts, USA. Struwe, L., S. Haag, E. Heiberg& J. R. Grant. 2009. Andean speciation and vicariance in Neotropical Macrocarpea (Gentianaceae-Helieae). Ann. Missouri Bot. Gard. 96: 450-469. Swofford, D. L. 1997. PAUP*. Phylogenetic analysis using parsimony, version 4.0. Sinauer Associates, Sunderland, Massachusetts, USA. Vogel, S. 1965. The role of scent glands in pollination. Smithsonian Institutional Libraries and National Science Foundation, Washington, D.C., USA. Whitten, W. M., M. Blanco, N. Williams, S. Koehler, G. Carnevali, R. Singer, L. Endara& K. Neubig. 2007. Molecular phylogenetics of Maxillaria and related genera (Orchidaceae: Cymbidieae) based upon combined molecular datasets. Amer. J. Bot. 94: 1860-1889. Young, K. R., C. Ulloa-Ulloa, J. L. Luteyn & S. Knapp, S. 2002. Plant evolution and endemism in Andean South America: an introduction. Bot. Rev. 68: 4-21. Zwickl, D. J. & D. M. Hillis. 2002. Increased taxon sampling greatly reduces phylogenetic error. Syst. Biol. 51: 588-598.
LANKESTERIANA 11(3): 253. 2011.THE GENUS CORYANTHES: A PARADIGM IN ECOLOGY GNTER GERLACH ABSTRACT. Coryanthes has fascinated naturalists, but other aspects of Coryanthes of 150 cm with the ant nest comprising 80 cm. Both organisms share a destiny because the plant is condemned to rapidly. Coryanthes Coryanthes family. Capsule ripening is also rapid; most species require only two months to mature a capsule containing type of insect can effect pollination. A bee that is too large for the passage between the epichile and column Coryanthes different scents, helping the taxonomist to delimit species that in nature are pollinated by the same bee in spite Coryanthes RESU M EN. Coryanthes fascin a los naturalistas, Coryanthes crecen diametro, solo el nido de las hormigas mide unos 80 cm. Los dos organismos son dependientes uno del otro, la planta esta condenada a la muerte si la colonia de hormigas asociada muere. Las plantas ofrecen nectar en del suelo. La alimentacin abundante permite a la planta un crecimiento muy rpido. Plantas de Coryanthes g, el cual es otro record dentro de la familia de las orqudeas. Las semillas para madurar necesitan solo 60 dias, una cpsula contiene alrededor de 600.000 granos de semilla, eso probablemente es una adaptacin a su
Las diferentes especies adentro del gnero Coryanthes por la misma especie de abeja. Todas estas peculariedades de las especies de Coryanthes menciondas aqu cabalidad en totalidad una serie de procesos. KEY WORDS : Orchidaceae, Coryanthes, ecology, pollination orchids, plants of Coryanthes were in great demand by horticulturalists. This was mainly because of their these plants a hard nut to crack. From the beginning story of Coryanthes and their biology started 30 years than 30 species in the hot and humid greenhouse, and me to work on this genus, because no one in the Neotropics could grow Coryanthes well for any length of time. The genus Coryanthes was established by Curtiss Botanical Magazine together with a colored plate. Sixty-two before 1980, so 27 species were described within the last 30 years. The scarcity of Coryanthes in herbaria frequent in the wild; 2) they are well protected by three days. These factors result in infrequent collection of Coryanthes by botanists. Plants of Coryanthes grow in humid lowland may simply represent a collection artefact caused by the easy accessibility from a dugout, the easiest way to Coryanthes picturata Coryanthes vasquezii Coryanthes macrantha Coryanthes speciosa espiritosantense Ruschi). Coryanthes species grow epiphytically in so-called ant-gardens (Fig. 1), which are the most complex and sophisticated of all mutualistic partnerships between of masses of soil, detritus, and chewed plant parts assembled at the branches of trees, forming huge clumps that may reach diameters of around 80-100 cm. All plant members in these partnerships are highly adapted, most of them obligate ant-garden plants. They are found in different plant families, e.g. Gesneriaceae (Codonanthe), Araceae (Anthurium), Cactaceae (Epiphyllum), Bromeliaceae (Aechmea), Piperaceae (Peperomia), Orchidaceae (Coryanthes, Epidendrum imatophyllum obligate; or arils that induce the ants to collect the seeds and place them in their nests. As the plants grow, nourished by the soil and other materials, their roots become part of the framework of the gardens. The ants in turn controlled by the ants; legitimate members are tended pruned by them. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.254 LANKESTERIANA
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. GERLACH The genus Coryanthes: a paradigm in ecology255 In Coryanthes ant-gardens, the ants belong mainly to the genera Crematogaster, Azteca (Myrmicinae), and Camponotus Coryanthes new shoots, bracts, and the outer surfaces of the Coryanthes species are Coryanthes orchids. The question in this mutualistic relationship is whether a Coryanthes existing ant nest or the Coryanthes that the Coryanthes seed germinates deep in an ant nest and that the young seedling with its long internodes reaches the surface of the nest and then change its seeds of an Epidendrum species growing in ant nests that lipid deposits within Coryanthes seeds may also the ant nest could be explainable but still needs to be hard to imagine that the wind-dispersed balloon-seeds liberated by Schleuderhaare (hairs in the capsule that help to disperse seeds when the capsule opens) are trees carry more than one ant-garden with Coryanthes (G. Bergold, T. Graf, and I) found an huge mango (Mangifera indica L., Anacardiaceae) tree with more than 10 Coryanthes Coryanthes plants to imagine that ants collect Coryanthes seeds from a capsule opening in one of their ant-gardens and carry circumstances. The genus Coryanthes sections, C. sect. Coryanthes C. sect. Eucoryanthes) with a smooth mesochile and C. sect. FIGURE association consists of Epidendrum, Epiphyllum (Cactaceae), Peperomia (Piperaceae), Anthurium (Araceae), and some Bromeliaceae. (Photo: G. Gerlach)
Lamellunguis with a mesochile bearing warts or lamellae. Micromorphology of the seeds (Plate 3) are all elongate, but those of C. sect. Coryanthes are fusiform, whereas those of C. sect. Lamellunguis only one tapered side. Seed ripening is remarkably quick in Coryanthes species; most capsules need et al., 2009). Similarly short times for seed ripening elsewhere in Stanhopeinae are found only in some species of Gongora, which occupy similar habitats. The other genera within this subtribe need more than six months to produce ripe seeds. well as a third clade composed of species with a nearly including the isolated species C. macrocorys Rolfe, the unclear. Erection of additional sections within the resolution and support. Coryanthes proterogynous. In Aristolochia L. (Aristolochiaceae) its pollen load at the stigma and cannot escape. On the releasing pollen onto the pollinator; the male and female phases are separated by time. Finally, the trap the exit of the pollinators. In Coryanthes, the anther; the separation of the sexes is here by space. Looking at the micromorphology of these trapLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.256 LANKESTERIANA PLA TE
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. GERLACH The genus Coryanthes: a paradigm in ecology257 FIGURE 2. A single most-parsimonious tree from cladistic analyses of nrITS sequences of Coryanthes (Whitten and Gerlach,
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.258 LANKESTERIANA wet from the liquid dripping from the basal glands and stored in the bucket. The liquid is not pure water (nor nectar) but contains some soap-like substance (saponins?) that disrupt surface tension of the liquid. The walls of the bucket formed by the lip show cell Cephalotus follicularis Labill. (Cephalotaceae) and Darlingtonia californica Torr. (Sarraceniaceae), which are not related to orchids (Barthlott and Ehler, 1977). PLA TE Coryanthes speciosa espiritosantense (Photos: G. Gerlach). PLA TE 3. SEM of the seeds of Coryanthes. From left to right, upper row C. sect. Lamellunguis: Foldats, C. trifoliata; lower row C. sect. Coryanthes: C. gernotii G.Gerlach & Romero, C. albertinae (Photos: G. Gerlach).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. GERLACH The genus Coryanthes: a paradigm in ecology259 bees from climbing out of the bucket. The column also forms part of the trap; the cells lie side by side and also Coryanthes are pollinated by male, perfumecollecting euglossine hidden under the mostly cup-shaped hypochile of the and reward for the bees. Each bee species has speciesgeographically and seasonally. In addition to orchid wood or feces. After collecting the fragrance compounds at the horn-shaped osmophore below the hood (hypochile) of the lip using its feathery, foretarsal brushes, the male with them, the pollinator comes into contact with drops on the pleuridia (liquid-secreting glands at the base of the From here the bee cannot escape because of the smooth surface of the inner side and its now-moistened wings. and directs the bee towards the exit passageway formed by the tip of the lip and the column apex. The claw of bit apart from the column, widening the exit so he can catching the stipes and freeing the pollinia from the exit channel, the pollinarium is glued to the rear of his any structures that he can grasp with feet or mandibles (Plate 4). After successfully exiting the passage formed PLA TE 4. Pollination of by Euglossa alleni in Costa Rica. From left to right: pollinator encompasses
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.260 LANKESTERIANA some time to dry itself. The entire pollination process lasts from 10 to 30 minutes depending on the presence or absence of the pollinarium. among sympatric species. Second, it reduces the and the orchid; if a bee too large for the exit canal is attracted and falls in the epichile, it cannot escape and pollination because it does not touch the stigma and pollinarium. G.Gerlach (among other species) collected by the headspace technique includes two interesting chemical dusty, leathery but still sweetish odor that is typical for C. mastersiana Lehm. (Gerlach & Schill, 1989; Kaiser, 2006). This species grows in the lowland forests and from 80-99%. Field studies carried out in Colombian Choc near Bahia Solano with the synthetic product Euglossa chalybeata and Euglossa asarophora. Ten FIGURE 3. GC/MS analysis of the trapped scent of : 1 Sabinene; 2 Eucalyptol; 3 trans-Sabinene trieno-5-lactone. Courtesy of R. Kaiser (2006).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. GERLACH The genus Coryanthes: a paradigm in ecology261 attractants. of chemicals in the fragrance of this Coryanthes species. This substance was found as a characteristic substance in the fragrance of C. albertinae Karsten in the habitat of that species with this synthetic alkene PLA TE 5. Variability within Coryanthes mastersiana G. Gerlach)
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.262 LANKESTERIANA PLA TE Trevoria glumacea; middle Gongora lagunae; left below -Stanhopea anfracta; right below Coryanthes trifoliata Pictures not to scale. (Photos: G. Gerlach)
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. GERLACH The genus Coryanthes: a paradigm in ecology263 green apple, characteristic of that chemical, but not a single euglossine bee was attracted. Two other baits on the same morning attracted 12 bees carrying pollinaria of C. albertinae. The failure was explained 10 years later when an oxygenated compound with a much longer retention time was found by Kaiser (2006) in the fragrance of C. albertinae bait was exposed to tropical rain or full sunlight. This By contrast, eucalyptol (2), used by most researchers only about an hour, depending on the air temperature. patterns may occur in different genera. Because of plants may attract the same euglossine bee species as pollinators without the risk of producing intergeneric the examples below, mechanical isolating mechanisms (pollinarium shape and placement) are critical, but related taxa. Ipsdienol with its precursor myrcene (see also Whitten et al., 1988) dominates the fragrance of Stanhopea anfracta Rolfe (40%/25%; ipsdienol/ myrcene ratio of total composition), butcheri Gongora lagunae G.Gerlach (40%/33%), Trevoria glumacea Garay (49%/16%), and Coryanthes trifoliata C.Schweinf. (51%/32%; Plate 6). These species place pollinaria at different sites on the bees: legs (); legs of presumably the right-hand side (Trevoria the abdomen (Gongora, Stanhopea); and below the scutellum but directed toward the head (Coryanthes). Similarly, methyl salicylate (wintergreen oil) occurs as large percentages in fragrances of Houlletia lowiana (84-98%), Stanhopea candida (55%), and Coryanthes leucocorys pollinia attached in different locations or orientations on the bees. interesting aspects of the ecology of Coryanthes. The different species are highly adapted to their habitats also be able to enjoy the study of Coryanthes in the wild. ACKNOWLEDGMENTS My thanks go to Gerd Seeger, who encouraged me to work on this genus and established a good base with the collection of Coryanthes in the Botanical fragrance analyses; Mark Whitten, who carried out the molecular work, discussed Coryanthes, on the text; Bert Klein and his team of gardeners in the plant (Synthesis of Systematic Resources) and the European Union for an Integrated herbarium research. LITERA TURE CITED Richter. 2009. Published on the Internet; http://www. orchidsrepbiol.de/cgi-bin/RBO/seed_ripening/display. pl/ accessed 2009. Barthlott, W& N. Ehler. 1977. Raster-Elektronenmikroskopie 19: 36. suggestions for future inquiries. Pp. 155-171 in: E. Physiological ecology of plants of the wet tropics. W. associations with ants. Pp. 200-233 in: U. Lttge (ed.). Vascular plants as epiphytes: evolution and ecophysiology. Springer, New York. Mesoamericanae I. Lankesteriana 8: 23-30. Gerlach, G. & R. Schill. 1989. Fragrance analyses, an aid to taxonomic relationships of the genus Coryanthes 168: 159-165. Gerlach, G. & R. Schill. 1993. Composition of orchid scents attracting euglossine bees. Bot. Acta 104: 379-391.
Coryanthes unter besonderer Bercksichtigung der Bltenduftstoffe. 83: 1-205. Coryanthes maculata 58: t. 3102. Kaiser, R. 2006. Meaningful scents around the world. Meyers, T. C. and Lamb, C. E. Published on the php?about=Coryanthes-Seedlings/ accessed 2009. 27: 2759-2760. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.264 LANKESTERIANA
Magazine published an article (Risen, 2008) related to the rights of plants under the new Constitution of forefront of international ecopolitics when it became nature, which includes the right to the maintenance and further, arguably granting broad protections to simple In order to comprehend the challenges we face within the new constitution should be understood. The of contamination and in harmony with nature. The underground resources otherwise known as the restoration of degraded natural areas is declared to be of public interest. nature has under this new Constitution, where Article 71 states the following: where life is generated and reproduced in nature, its existence has to be respected LANKESTERIANA 11(3): 265. 2011.WHAT WILL BE LEFT OF THE PRIMARY FORESTS IN ECUADOR? ALEXANDER HIRTZ ABSTRACT. Ecuador is among the smallest countries of South America (250,000 square kilometers) but has the highest density of human population with the highest network of primary and secondary roads, which also explains why Ecuador has the highest slash-and-burn ratio per capita in America (about 250,000 hectares per approximate 1000 endemic orchid species described to date for Ecuador are probably not growing in these RESU M EN. Ecuador es uno de los pases ms pequeos de Amrica del Sur (250,000 kilmetros cuadrados) pero que tiene la ms alta densidad de poblacin humana con la red ms alta de caminos primarios y secundarios, lo cual explica porqu Ecuador tiene la relacin ms alta per capita de corte y quema en Amrica y nunca ha sido estudiadas botnicamente. As, un gran porcentaje de las aproximadamente 1000 especies endmicas de orqudeas descritas hasta la fecha para el Ecuador, probablemente no estn creciendo en estas KEY WORDS
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.266 LANKESTERIANA Any person or community may make demands to for its restoration. They are also responsible to pay the damage. Articles 73 and 396 obligate the State to take the precautionary measures or restrictions to all the destruction of ecosystems or the permanent article in the Constitution because it implies that people and communities may, for example, collect This application of the article is a total contradiction to the current laws and regulations established by the impact, mitigate and restore any damages caused by management program. Legal actions to prosecute and Article 397 establishes the following: in case of restoration of health to the damaged ecosystem. The State will sanction not only the party responsible for damaging the ecosystem, but it will also make the may exercise legal actions against parties who might endanger the health of any ecosystem, which includes water, air, soil, and the life forms within them. The landholdings in fragile ecosystems, which include the pramos, wetlands, cloud forests, tropical dry and wet is considering reducing landholdings to between 5,000 and 10,000 hectares, in Ecuador the new legislation is considering limiting the landholdings to 800 hectares. which will raise questions from landowners who want management of all water resources, water basins, and the State will adopt adequate measures to mitigate the effects of global climate change by limiting gas emissions, deforestation, and air contamination. It will Constitution, all primary forests are protected, and any immediately, where the authorities may not delay any action described in the new Constitution by claiming that the bylaws and regulations are not yet in place that about 300,000 hectares of primary ecosystems State immediately restore any destruction and punish the culprits. In his article published by the Magazine how, exactly, a country as poor as Ecuador can protect these rights and how it can protect Pachamama or a surface of only 283,561 km2, Ecuador is second smallest among the countries of South America, after Uruguay. Ecuador has the highest density of human population in South America with 49 inhabitants per
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. HIRTZ What will be left of the primary forest in Ecuador?267 country four times larger, has only 8.3 inhabitants per country and another three million working abroad. and Ecuador has to assume that, of the three million next two years because they lost their jobs in the global economic crisis. Ecuador has the highest network of primary and secondary roads in South America, adding up to at least 40,000 kilometers. Ecuador is only 700 kilometers long, north to south, from the border of Colombia to the border of Peru. The growing capital These area constraints and population density factors are the main reasons why Ecuador has the highest ratio of slash-and-burn to total surface area of any country in the Americas; we lose around 300,000 hectares of primary forests per year. Because of the high density of plants growing on trees, about 20 to 30 of our generation seeing a dramatic decrease in the new constitution where nature has almost unlimited rights is a futile effort, because it will only slow down and oil companies and rarely address the real problem, by the poor. The efforts of a handful of concerned botanists trying to stop the ancestral practice of slashand-burn agriculture will not change the fact that it is an enormously successful adaptation to the rigors and constraints of the tropical forest. within which at least two thirds of the areas are Unfortunately, a paragraph in Article 405 of the new Constitution states that neither a foreigner nor any concessions in protected areas. This will undoubtedly raise legal issues in cases where foreigners or NGOs as a protected area, only to lose it under the new Constitution. A large percentage of the approximate 1,000 endemic orchid species described to date for Ecuador are probably not growing in the currently established to the road. The remaining 80% of land in Ecuador Constitution, probably into areas no larger than 500primary forest left had better replace it with pasture, as was customary in the 1960s and 1970s under the Agrarian Reform. Of course, this threat directly contradicts the new Constitution. It can be concluded that no primary forests become extinct in nature. To make matters worse, loggers will not impact the protected areas in the oncoming climate changes, regional acid rains, and cannot ask for more than what is already written in the new Constitution. But it is clear that the other forests or much longer. Therefore, implementation of botanical gardens distributed at all or orchid photo safaris is one of the main methods to learn that this will bring business and well-being to
public places, particularly orchids and bromeliads that they rescued from fallen trees. The Ministry of Bancos are good examples. This year the Orchid Society of Quito has decided was proposed and will be managed by a network of Napo that are currently forming the Mushuk Sisa Foundation. These communities are already well tourism. Their botanical garden, highlighting orchids, will be established next to the Ro Jatun Yacu. We of the Quito Orchid Society are dedicated and diligence we will accept the challenge and succeed It is our hope that other orchid societies both here in challenge and help sponsor small communities in their nascent botanical garden projects. These communities are our keys to success and the future of orchid tourism and orchid LITERA TURE CITED LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.268 LANKESTERIANA
et al., 2009). Increase in species richness from the poles to the tropics (Pianka, 1966; Rohde, 1992; Willig et al 1995) is still one of the main topics in contemporary important determinant of species richness (Wright, 1983; Wylie & Curie, 1993a,b; Pelkey et al., 2000; LANKESTERIANA 11(3): 269. 2011.HOW UNIFORM IS SPECIES DIVERSITY IN TROPICAL FORESTS? PA VEL KINDLMANN 1 & CARLOS A. VERGARA CASSAS 21 2 ABSTRACT. the region. The sampling effort can also account for the fact that most endemic orchid species are found close RESU M EN. Uno de los factores que genera preguntas es la distribucin de las especies, misma que se podra ser un resultado ms propicios para especies terrestres, mientras que los que se encuentran al interior del bosque son apropiados par KEY WORDS :
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.270 LANKESTERIANA et al., 2005; Storch et al., 2005). Area is clearly also be important. One of these factors is the sampling effort the the number of orchidologists in the region. The plant species richness in areas where botanists are disproportionately present as an artefactual consequence of a more thorough sampling (Pautasso & McKinney, 2007). For orchids this is illustrated in Figure 1 using the are similar to each other in many respects: they are of habitats and similar climatic conditions. Ecuador regression line. There is much less known about orchid The sampling effort can also cause the effect et al. (2007) most endemic orchid species were found close to the roads. Thus it Description of study area of the Uchumachi mountain close to the settlement the Andes. yungas secondary forest, and primary forest. The secondary ferns (Cyathea amaznica Inga sp.), walnut (Juglans boliviana (Cecropia angustifolia FIGURE that for temperate ones. The R2 FIGURE 2. The approximate position of the study area is indicated here by the red dot. FIGURE 3. The the study area.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KINDLMANN & VERGARA CASSAS 271 of taller trees, wider in diameter, and a reduced understory due to a reduction of light in the forest forest is dominated by tree ferns (Cyathea amaznica), Clusia haughtii Sapium aereum Ficus obtusifolia Roxb.) for a large part of the basal area. Other species such Miconia guianensis Cogn.), wild papaya ( which indicates that they may be under the threat of extinction (Endara, 2001). data from an on-site weather station shows that the July) about 15 C, with maximum yearly temperatures around 25 C and minimum yearly temperatures of 15 C. The total annual precipitation is 2390 mm, with 300 mm per month) and no month with less than 80 Methods We used linear transects perpendicular to the road depicted in Figure 3. Along the transects, we set up species were determined. The accuracy of species plots, 33 were adjacent to the road (habitat R), 40 were in the secondary forest (habitat S), and 44 were in the primary forest (habitat P; see Fig. 4). Results exponential pattern, when the species were ranked from the commonest to the rarest (Fig. 5). Table 1 shows the (RSS) of the function y = a.e between the number of squares in which the species was found and species rank, when the species are ranked from function y = a.e FIGURE 4. Plan of the study area. The thick black line is the road. Thin lines indicate the transects, and small circles are the plots. FIGURE 5. Number of squares in which the species was found against the species rank, when the species are ranked from the commonest to the rarest.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.272 LANKESTERIANA when the species are ranked from the commonest to the epiphytic and 18 terrestrial species), only one species occurred in all three habitats, the epiphytic Pleurothallis Rchb.f. There were 3 epiphytic and 6 terrestrial species, which occurred in two habitats. The remaining 9 epiphytic and 12 terrestrial species were square that were found at the edge of the road, in the secondary forest, and in the primary forest for terrestrial and epiphytic species. Clearly, both the number of of terrestrial species strongly decreased toward the An opposite situation occurred in epiphytic species. Both the number of epiphytic species and the number toward the interior of the forest. The smallest number followed by the secondary forest, and the primary forest with the largest number of epiphytic species and Discussion and conclusions We found opposite trends in terrestrial and TABLA 1 squares (RSS) of the function y = a.e relationship between the number of squares in which the species was found and species rank, when the species are those of the function y = a.e species rank, when the species are ranked from the TABLA 2 forest, P primary forest). Species common to all three habitats is in red and highlighted in yellow; species common to two habitats are highlighted in blue.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KINDLMANN & VERGARA CASSAS 273 epiphytic species increased toward the forest interior. did the total number of species and the total number when making conclusions about the trends in orchid mode should be taken into account. The reasons for the interior, density of the trees increases, and therefore Therefore, habitats close to the openings (roads, species, whereas those deep in forest interior are more host trees. ACKNO W LEDG M ENTS The research was supported by the LITER A TURE CITED Arrhenius, O. 1921. Species and area. J. Ecol. 9: 95-99. Endara, A. R. 2001. Inventario de las especies forestales del bosque hmedo tropical premontano del Cerro Uchumachi sector Carmen Pampa. Patrones de endemismo de orqudeas endmicas in: A. M. Pridgeon & J. P. conference on Andean orchids Particular de Loja, Loja, Ecuador. species-energy relationships. J. Anim. Ecol 74: 498507. area. Ecology 3: 158-162. Morales, C. B. 2004. Manual de ecologa. 2. edicion. Pautasso, M. & M. L. McKinney. 2007. The botanist effect Biol. 21: 1333-1340. Pelkey, N. W., C. J. Stoner & T. M. Caro. 2000. Vegetation 297-309. the search for the primary cause. Oikos 65: 514-527. 80: 172176. 142: 23292334. area-energy relationship. Ecol. Letters 8: 487-492. Evaluacin de la erosin hidrica en un (Coffea arabica) (Inga adenophylla) chonta y machete en Carmen Pampa. Tesis de grado. FIGURE 6 were found at the edge of the road, in the secondary forest, and in the primary forest for terrestrial and epiphytic species.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.274 LANKESTERIANA in: A. A. Myers and P. S. Giller (eds.). Analytical biogeography. species-area theory. Oikos 41: 496-506. and patterns of species richness: I. Patterns of bird, angiosperm, and mammal species richness on islands. Biol. Conserv. 63: 137-144. Wylie, J. L. & D. J. Currie. 1993b. Species-energy theory and patterns of species richness: II. Predicting mammal species richness on isolated nature reserves. Biol. Conserv. 63: 145-148.
Charles Darwin geologist is noteworthy to commemorate his seminal research relating earthquakes and mountain building to America. The purpose of this paper is to apply and reasons why southeastern Central is such a hotspot LANKESTERIANA 11(3): 275. 2011.ACTIVE MOUNTAIN BUILDING AND THE DISTRIBUTION OF CORE MAXILLARIINAE SPECIES IN TROPICAL MEXICO AND CENTRAL AMERICA STEPHEN H. KIRBY ABSTRACT. molecular phylogenetics, the Camaridium clade is easily the most prominent genus in Central America and is largely restricted to the highlands of Costa Rica and Panama, indicating that this region is the ancestral home of this genus and that its dispersal limits are drier, lowland cordilleran gaps. The mountains of Costa Rica and southeastern Central America. RESUMEN. aparece que el caldo Camaridium es el gnero ms prominente en Amrica Central, y est restringido a las tierras altas de Costa Rica y Panam, indicando adems que esta regin es su lugar de origen y que sus lmites para su dispersin son las tierras bajas y ms secas. Las montaas de Costa Rica y Panam se encuentran entre de numerosas placas tectnicas. Considerando la informacin geolgica disponible, se concluye que el rpido orqudeas en el sur-este de Amrica Central. KEY WORDS : Central America, geology, geography, Orchidaceae, Maxillariinae, distribution
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.276 LANKESTERIANA interplay between mountain belts (and associated lowland gaps), geologic processes, and biological foremost a naturalist and that early in his life as a scientist he made important contributions to geology. (1809-1882), a freshly minted graduate of Cambridge HMS Beagle 1875) famous series, The Principles of Geology (1830, 1832, 1833), a landmark treatise about the rock record Sedgewick (1785-1873) in late summer of 1831. With this limited knowledge and experience in geology, but to create conceptual models of how nature works, he explored the globe with his shipmates. In the decade after the conclusion of the Beagles that became founding documents in the 19th century 1838, 1839, 1842, 1844, 1846). In three of these the Beagles First, he documented the numerous formerly marine water marine seashells that were indistinguishable Beagle at anchor et al., 1997). At about 11:40 a.m. local time, a great subduction earthquake occurred ground motion lasted two minutes; the near-shore area of the town was damaged due to the seismic sea Beagle was hit with jarring motions as if it had run aground (Yeats et al. large region of coastal uplift that was produced by the earthquake. Although these uplifts are sudden during earthquakes, such shocks occur infrequently and thus led a mule-pack-train exploration of the high Andes to the passes between present-day Santiago and Portillo the foot of Aconcagua, the highest peak in the Andes. that the earthquakes somehow represented the motions that raised the shorelines, produced the raised marine terraces, and slowly built the Andes mountain range all together. It would be decades later before it was generally understood that most tectonic cordilleras like deformation or shortening, leading to thrust-faulting and consequent thickening (e.g., Fisher, 1881; Suess, 18831909), and that continuous belts of earthquakes are often coincident with or parallel to such mountains (Mallet, 1858; Milne, 1886). It was more than a deformation of continental margins is generally a consequence of subduction motion of oceanic plates as and Isacks, 1967; Isacks et al
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution277mountain building: near where tectonic mountains are being built. In the context of modern digital seismology, earthquakes can be monitored anywhere in the world using global stations, complete down to about Richter magnitude mountain building is occurring at present. Likewise, mountain building, although generally slow by human standards, can change the distribution of climatic indicated to him that this mountain range represented early work as an insightful pioneer in geology also has a direct bearing on our understanding of one of 1859, 1862; Wallace, 1858, 1870, 1889). This paper is a preliminary exploration of these concepts as applied to southeast Mexico and Central Origin of Species. It is intended as a preliminary application of shaped in part the distribution of orchid species and their Homo, in Africa. Southeast Central America: a hotspot of orchid diversity The status of southeast Central America as a et al., 2000; Ossenbach, 2009). Although there are strong differences in the likely degree of undersampling of loss by country, the data from a recent checklist of orchid species distribution in Central America and southeast Mexico show that differences in total orchid 1A, B) are extremely large (Ossenbach et al., 2007), probably more than expected due to differences in the degree of undersampling. In particular, the species area density contrasts with the rest of tropical America are usually not natural biogeographic boundaries, but orchid species distributions are often known only to the are stark differences between the large area densities of endemic orchid species of Costa Rica and Panama FIGURE 1 geographic area (data source: Ossenbach et al. (2007)). A. Total species and endemic species numbers by country. B. Species per 1000 km2 country area (species density) for both total species and endemic species.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.278 LANKESTERIANAone hand and the low species densities of the largely transcend sampling biases. An important question is why such differences exist, an enigma that is the subject of this paper, one that builds on the work of southeast Central America. Mountain belts, geological processes, and biological diversity It is well established that wet tropical premontane and biologically exuberant regions in the world (e.g., regional climatic conditions through the orographic upslope that increase rainfall and also the effects of and promoting plant growth. The mountain belts in of moist premontane to montane forests from southern Mexico to Panama (Fig. 2). I discuss below other effects of mountain chains (and processes that occur in them) on the biogeography of tropical orchids within the Americas in the context dispersal, speciation, and extinction (Fig. 3). I consider these processes in this context of the ensemble of orchid species and their biological cohorts FIGURE 2 Map showing the eco-regions of Central America and southeast Mexico with emphasis on moist, broadleaf, et al. (2001)].
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution279enable them to germinate. It should be understood that any biogeographic conclusions reached in this report pertain to the ensemble of these cohorts. The spectrum of active geological processes, environmental change, and evolution: dispersal, extinction, and speciation Dispersal South America. Mountains in the tropics represent lowland conditions, because lowland tropical species are not forced to adapt to large seasonal temperature i.e., in the tropics, montane and lowland temperatures dispersal of orchid species adapted to cool, moist to occurrence of orchid species on Cocos Island, some 530 km from mainland Costa Rica, some of which are endemic to the island and some found elsewhere in Central America and Peru (Trusty and Blanco, 2005; Trusty et al., 2006). The major lowlands that interrupt southeast Mexican and Central American cordilleras are from north to south (Fig. 4): 1) the Tehuantepec (Chiapas/Oaxaca) gap; 2) the Nicaraguan depression; 3) the GatunBalboa gap crossed by the Panama Canal; 4) the and Caribbean coast ranges in eastern Panama; 5) the R.o Chucuaque/Gulf of San Miguel gap; and 6) the near-coastal Colombian lowland between the northwest Colombia from the main Andean cordillera (the Cordillera Oriental). The Nicaraguan depression is easily the most prominent lowland gap among mountain ranges in Central America based on its width and the fact that some of this lowland is occupied by the lowland Lakes Nicaragua and Managua and the depression is a rift structure associated with extension and subsidence (Phipps Morgan et al., 2008; Funk et al. broad inland seaway that connected these bodies of water existed as recently at 6,000 years before present (Roberto Protti, personal communication, January mountains can be many hundreds of meters lower in Volcanic Range in Costa Rica. It is not known if these FIGURE 3 Schematic ternary diagram depicting graphically biota (physical and biological changes) and the processes that control the biogeography of life forms: dispersal, speciation, and extinction. See text for discussion. FIGURE 4 Color-shaded relief topography of southeast Mexico, Central America, and northwest Colombia showing the principal lowland gaps between cordilleras.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.280 LANKESTERIANA will be discussed after interpreting distributions of taxa of Maxillariinae by country. SpeciationSpeciation is dominantly a biological process but one that can factors. Mutagenesis, the production of new biological forms with inheritable traits, is caused by damage to the nucleotide sequence of organisms and is affected by exposure to chemicals, high-energy radiation from natural decay products of radiogenic nuclides in nature, isolation, can lead to new species that do not cross Gillespie, 1967). Mountain building clearly can cause It is often assumed that exposures to mutagens are random (and associated with normal biological change are essentially constant, and hence that branching of the tree of life can be accurately dated by Volcanoes produce a toxic brew of chemicals in gaseous interact in the atmosphere to produce sulfuric, carbonic, kill plants and create dangers to human health. It is not known if sub-lethal exposures of these chemicals can induce mutagenesis in plants, a question that could be such as uranium, thorium, potassium (K40), and radon. health risks with long exposure. These examples might indicate that mutagenesis may not be spatially random and could occur at higher rates in tropical highlands. This possibility should be experiments. Putting the question another way, such experiments would answer the interesting question: and faster in some mountain ranges? A s natural agents of destruction of habitats caused biotic extinctions of species endemic to those orchid dispersal in the geologic past. manifold effects on conditions for life on Earth (Mason et al., 2004) Such eruptions can launch columns of hot tephra (ash) and gas as high as 50 km into the stratosphere. Volcanic aerosols suspended high in the crises are well documented in ice cores drilled from ice sheets in Greenland and Antarctica. Sequences of such of time and hence had global climate impacts that are by large eruptions of magma associated with largeAmerica. Another geologic indicator of the scope of
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution281hot eruption plume, a phenomenon called base surge, narrowly endemic orchids in the past. Repopulation of Krakatau, the island in the Sunda Straight left after the cataclysmic 1883 eruption in Indonesia, now has a dense tropical forest in place (Simkin and Fiske, 1983). probably caused extinctions of narrow endemics in Volcanic gases that boil out of magmas as they ascend to the surface during eruptions interact with atmospheric water to make acid rain and VOG (a ground-hugging certain conditions during prolonged eruptions, VOG can drift hundreds of kilometers from its source taken place between orchid populations. Chronologies absolute time marks on molecular clocks. Young geological history of Costa Rica and Panama The mountains in Costa Rica and Panama are, along with those in western Colombia and certain cordilleras in Ecuador and Peru, among the youngest in tropical Latin America. This condition is largely a consequence Carnegie Ridges) that originate at the Galapagos hotspot of Costa Rica, Panama, Colombia, and Ecuador (Fig. 5; Mann and Corrigan, 1990; Mann, 1995; Coates, 1997; et al. Mann et al., 2006; Sak et al. documented the remarkably young ages of the major phases of mountain building of these cordilleras (Table 1, Fig. 6), ranging from the Guanacaste and Central Volcanic Ranges (< 0.5 Ma BP [million years before << 2 Ma BP, 5 Ma BP for Cordillera Tilarn, and 4-10 Ma BP for the main Talamanca Range that extends into Panama and forms the mountainous backbone of both countries (sparsely distributed older igneous rocks in the Talamanca may represent the roots of earlier island-arc consequence of underthrusting in the former forearc basin rocks of the Fila Costea under the Talamanca, a process that started no earlier than 2 Ma BP (Fisher et al., 2004; Morell et al., 2007; Steichler et al., 2007; Sak et al., 2009). Thus the highest mountain range in Costa Rica and Panama was probably uplifted to its These cordilleras are among the youngest mountain ranges in the world and were largely built during the time when our own species in the genus Homo Costa Rica and Panama are the youngest products of continent and mountain building in tropical Latin America. This region was formerly an oceanic seaway earlier dispersal of plant and animal species, famous in the annals of biogeography for the later great faunal exchange that was in full force by the beginning of Quaternary time (2.6 Ma BP; Webb, 1997). The gradual closing of this seaway by sediment accumulation and mountain building between present-day Colombia and the rest of Central America began about 12 Ma BP and was completed about 3 to 4.2 Ma BP, based mainly a chronology of ocean circulation through the seaway
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.282 LANKESTERIANAbetween the Atlantic-Ocean/Caribbean-Sea and the et al., 2001). Figure 2 shows the distribution of moist, tropical, broadleaf montane forests in Central America based the windward fronts of mountain ranges, based on northeast margins of moist, broadleaf montane forests in Central America and tropical Mexico. This map clearly shows that the total lengths and areas of such eco-regions in Mexico exceed those in Costa Rica part of tropical Mexico are far older that those in similarities to those of southern Mexico. Geologically represent rocks accreted to the Central American et al. these highlands are isolated mountains representing rock types that are resistant to erosion; many of these uplands are dominated by Pinus species. The presentNicaragua support mostly lowland forest. Although forests are in lowlands. Finally, the Central American isthmus is narrowest in Costa Rica and Panama, and orographic effects of mountains tend to distribute FIGURE 5 Plate-tectonics map of tropical America, including plate names and plate boundaries. Also shown are locations et al. (2006).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution283 isthmus than farther north. alone does not explain why so many more orchid species are endemic to the southernmost countries be most exuberant. Of course, the latter are at lower latitude than tropical regions farther north, and perhaps at higher latitudes. For many orchid tribes, centers day orchid species distribution may be a snapshot of a general trend toward northward dispersal from two working hypotheses in this study: 1) hotter and drier lowland gaps between the cordilleras of Central America reduce the rates of this northward dispersal of species adapted to highland conditions (and possibly to northern Central America), and 2) high rates of in promoting more rapid orchid speciation in Costa Rica and Panama. These biologically important factors associated with mountain building are applied to the country distribution data for core Maxillariinae in Central America and southern Mexico. A case study: distribution of Maxillariinae in Southeast Mexico and Central America Data and methodology distribution of species in core Maxillariinae in Central good taxa to use. There is a large number of species (550580) according to Whitten et al. (2007) and display a range of pollination syndromes. Ossenbach et al. (2007) documented 160 species in Central America and tropical Mexico (the states of Chiapas, Tabasco, found in all of the countries of South America and Whitten et al. (2007) using multiple molecular markers, resulting in a well-supported generic realignment of the subtribe into 17 clades (Table 1). This was followed up et al. (2007) based on the molecular data and phenotypical characters described by Whitten et al. (2007). TABLE 1 Core Maxillariinae cladogram (Whitten et al., 2007). Estimated total species numbers and established species counts in each genus from Blanco et al. (2007). FIGURE 6 Color-shaded relief map of Costa Rica showing the locations of the principal mountain ranges, their et al October 2009).
My starting point for country distribution data on species in core Maxillariinae are the checklists by Ossenbach et al. (2007) and Atwood (2003). I supplemented these resources with the following online herbarium data resources: Tropicos (Missouri Botanical Garden), INBIO, Lankester Botanical Gardens Epidendra and the To assessments in the online Tropicos checklist and papers by Atwood and Mora de Retana (1999) on the subtribe. For Central American nations, I also relied on the most recent orchid monographs by countries listed below in Literature Cited. For tropical Mexico, I also consulted Soto et al. et al. (2005). In the interest of exploring distribution differences between Central and South America, I checked the distribution of species found in Central America and tropical Mexico Antilles by consulting Misas-Urreta (2005) for nearcoastal northwest Colombia; an unpublished database species Native Colombian Orchids edited by Escobar (1990) and for Peru; McLeish et al. the Antilles. In cases of differences in distribution data between sources, my preference was generally to adopt Many ambiguities exist in this distribution database, data for specimen collection were generally taken as those reported except where geographic locations were degrees of undersampling. Of particular concern is undersampling in northwest Colombia and eastern Panama due to security challenges, a shortcoming that may bias assessments of the distribution data of species common to both Central and South America. Panama (Pa), Costa Rica (CR), Nicaragua (N), El (Be), Mexico (Mx), Greater Antilles (GA, including southern Florida, USA), Lesser Antilles (LA), Guyana Results Geographic distributionThe country distribution (Table 2): Class 1: Species endemic to Costa Rica and/or Panama (Southeast Central America SE CAm) [83 species in this class] Class 2: Species endemic to both SE CAm and at least two countries in northern Central America (N Class 3: Species endemic to both SE CAm and Ecuador, Peru) [31 species] Class 4. Species endemic to N CAm [9 species] Class 5: Pan Latin America (N CAm and SE CAm and NW S Am) [18 species] Class 6: Occurrence in Ossenbach et al. (2007) not in SAm [6 species]. TABLE 2 core Maxillarinae species reported in Central America according to their north-south geographic spread by LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.284 LANKESTERIANA
endemic species among Central American nations et al. (2007) study. Second, more species in Maxillariinae in Costa Rica America (31) than other countries in Central America and tropical Mexico (13). Combined with the fact that nearly three-quarters of all species in Maxillariinae are found only in South America indicates that the primary that continent, with a secondary center in Costa Rica and Panama and with lesser endemism in northern Central America, tropical Mexico, and the Antilles. It is also useful to examine the northern and southern geographic limits by country of species in this taxa for species that occur in Central America and tropical Mexico (Fig. 7). By far, the largest number of species follows from the high number of endemics in these number of Mexican endemics (Fig. 7A), the northern limits of tropical climate, and also the longer potential dispersal distance to the Greater Antilles and southern Elevational distribution I show the distribution of 8 and 9 for two of the six distribution classes studied. Class 1 species (83 total), those with the narrowest mostly adapted to the cooler and wetter conditions found in the cloud-forest highlands of Costa Rica and Panama (Fig. 8A, 9A). There are nine exceptions to this trend: species that are restricted to Costa Rica and/ below 200 m (Camaridium suaveolens (Barringer) M.A.Blanco (Ames & C.Schweinf.) M.A.Blanco, Mormolyca dressleriana endresii Rchb.f., Ornithidium nicaraguense Garay) M.A.Blanco & Ojeda (Schltr.) M.A. Camaridium latifolium Schltr. (Rolfe) and Camaridium vittariifolium (L.O.Williams) M.A.Blanco according to the realignments by Whitten et al. (2007) and Blanco et al. (2007). Finally, no species in this geographic FIGURE 7 in Central America. A. Northern limits by country. B. Southern limits by country. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution285
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.286 LANKESTERIANA FIGURE 8 FIGURE 9 geographic distributions. A. Class 1 species (Costa Rica and/or Panama endemics), largely upland species with exceptions origin. For Class 5 species, the Pan Latin American and most (17) below 200 m (Fig. 8B, 9B). The wide geographic distribution of this class indicates that part of their successful dispersal may be rooted in their gaps between cordilleras. Geographic distribution of clades the distribution of the clades of Whitten et al. (2007) this study. Summing the species numbers for classes 1 through 5 shows that Camaridium is easily the most abundant genus of core Maxillariinae in Central America, representing 62 of the total of 72 species that Blanco et al. This important genus of tropical epiphytes represents about 15% of all species in the core Maxillariinae and
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution287 such as C. biolleyi (Schltr.) Schltr., C. bradeorum Schltr., and C. inauditum (Rchb.f.) M.A.Blanco to the striped miniatures Schltr. and C. tigrinum (C.Schweinf.) M.A.Blanco to the challenging C. cucullatum (Lindl.) M.A.Blanco complex. About 72% of Camaridium species are endemic to Costa Rica and with South America and three with the rest of Central America. Only three Camaridium species are reported to be endemic to other Central American countries, and It is therefore a reasonable inference that this genus Since species in this genus largely occur in highland Pliocene and Quaternary or the last 5 million years to 500,000 years depending on the mountain range). Only limited dispersal of this genus to other Latin American Except for two genera with a small number of species (Inti and Trigonidium), Central American endemic species in other genera in core Maxillariinae, et al. (2007) and listed in Blanco et al. (2007), represent minority populations compared to species in those genera in South America (Table 3). Notable among these genera are stricto (only 18% occur in Central America out of a total of 165 species placed in that genus by Blanco et al. (2007), the largest clade in the subtribe), (32% of 50 species), Mormolyca (16% of 25 species), (31% of 13 species), Sauvetrea (8% of 13 species), and Ornithidium (17% of 60 species). Since the geographic centroids of species in these genera are clearly in South America, it is plausible that they in Central America may be partly a consequence of the smaller land areas of Central American countries compared to South America. The balance of the 17 genera of Whitten et al. The genus Ornithidium has an estimated 60 species, Ornithidium species TABLE 3 Geographic distribution of species in core Maxillariinae following Whitten et al. (2007) and Blanco et al. (2007). See notes in text and at bottom of this table.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.288 LANKESTERIANAreported in Central America, four are highland species (O. hagsaterianum (Soto Arenas) Senghas, O. conduplicatum Ames & C.Schweinf., O. pittieri Ames, and O. repens (L.O.Williams) M.A.Blanco & Ojeda), extending down to lowland forests (O. nicaraguense O. adendrobium (Rchb.f.) M.A.Blanco & Ojeda, and O. fulgens Rchb.f.). Ornithidium fulgens, reported from hummingbirds (Fogden and Fogden, 2006) and is the most widespread of the genus. A number of species in this clade resemble O. fulgens (small, globose or reported to produce nectar). These species include: O. aggregatum Rchb.f., O. aureum Poepp. & Endl., O. canarense (J.T.Atwood) M.A.Blanco & Ojeda, O. coccineum (Jacq.) Salisb. ex R.Br. (the type species for the genus Ornithidium), O. conduplicatum Ames & C.Schweinf., O. giganteum Lindl., O. jamesonii Rchb.f., O. miniatum Lindl., O. pittieri Ames, O. ruberrimum (Lindl.) Rchb.f., and O. semiscabrum Lindl. in highland forests where hummingbirds are said to O. fulgens is spite of the presence of many lowland gaps in which Discussion data in this study. More distribution information is now Information Systems (GIS) to plot distributions on map such as the SRTM data (Shuttle Radar Topographic Mission). A recent GIS study by Lorena Endara of orchid distribution in Ecuador shows the power of such methods (Endara et al., 2009). Such methods may names) often recorded in older reports. Also, climate with orchid species distribution data. Some botanical institutions are already employing these methods. collections are often conducted in the campaign mode, i.e., go out and collect for a few days or weeks and undersampling because of possibly collecting outside actually sampled. The writer is co-founder of the in its 7th year. Orchid collections from downed trees and tree limbs are made during maintenance of such consuming, and rare. Finally, the density of sampling for molecular phytogenetics has generally not progressed down to in establishing clues as to dispersion pathways or geographic separations. As orchid genotyping becomes per species are sequenced, subtle differences in genetic species distributions alone. Conclusions Quammen (2007), author of the immensely popular and readable book about island animal biogeography, the Song of the Dodo that just as Copernicus (1473-1543) put the sun, rather than planet Earth, at the center of our solar system, Homo sapiens as just a mammalian species in a long succession of
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution289life forms shaped by natural selection. One can take this concept another step. As a geologist in an era of great strides in increasing understanding of our geography, and our climate into a chronology of a long on Earth that enables natural selection to work, but ACKNOWLEDGMENTS. family (Federico, Vanessa, and Federico, Jr., owners of I am especially indebted to Carlos Ossenbach for his encouragement and access to the orchid species database for the Central American Isthmus before publication, and been helpful in directing me to core Maxillariinae data and thank Alec Pridgeon for the opportunity to speak at the LITERA TURE CITED The Marie Selby Botanical Gardens illustrated dictionary of orchid genera. Cornell Flora Costaricensis, Family #39 Orchidaceae: tribe Maxillarieae: subtribes Maxillariinae and Oncidiinae. Fieldiana, Bot., n.s. 40. Atwood, J. T. 2003. Pp. 291-332 in: R. L. Manual de plantas de Costa Rica. Volume 3. Missouri Botanical Garden Press, St. Louis, Missouri, USA. Baxter, P. J. 2000. Impacts of eruptions on human health. Pp. 1035-1043 in Sigurdsson (ed.). The encyclopedia of volcanoes. Academic Press, New York, USA. L. Endara. 2007. Generic realignments in Maxillariinae (Orchidceae). Lankesteriana 7: 515-537. Burger, W. A. 1985. Why are there so many kinds of in: W. The botany and natural history of Panama. Missouri Botanical Garden Press, St. Louis, Missouri, USA. Coates, A. G. 1997. The forging of Central America. Pp. 1-37 in:. A. G. Coates (ed.). Central America: a natural and cultural history Connecticut, USA. Coats, R. R. 1962. Magma type and crustal structure in the Amer. Geophys. Un. Monogr. 6: 92-109. go.cr/herbario phaenomena, and on the formation of mountain-chains Proc. Geol. Soc. London 2: 654-660. Journal of researches into the geology and natural history of the various countries visited by H.M.S. Beagle UK. The structure and distribution of coral reefs. Smith, Elder & Co, London, UK. Geological observations on the volcanic islands visited during the voyage of H.M.S. Beagle. Smith, Elder & Co, London, UK. Geological observations on South America. Smith, Elder, and Co., London, UK. 1-52. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life 1st edition. John Murray, London, UK. On the various contrivances by which British and foreign orchids are fertilized by insects. 1st edition. John Murray, London, UK. Mapa geolgico de Costa Rica 2007 Librera Francesca S.A. Atlas tectnico de Costa Rica San Jos, Costa Rica. global tectonics. J. Geoph ys. Res. 75: 2625-2647.
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LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. KIRBY Mountain building and Maxillariinae distribution291Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. 403: 853-858. Nir, M. A. 2000. Orchidaceae Antillanae New York, USA. anomalous structures in the upper mantle and the lithosphere. J. Geophys. Res. 72: 4259-4275. Ossenbach, C. 2009. Orchids and orchidology in Central America: 500 years of history. Lankesteriana 9: 1-268. Orqudeas del istmo Centroamericano: catalogo y estado de conservacin (Orchids of the Central American ). Editorial 25 de Mayo, Sabanilla, Costa Rica. Phipps Morgan, J., C. Ranero & P. Vannucci. 2008. Intraarc extension in Central America: links between plate Planet. Sc. Lett. 272: 365-371. The reluctant Mr. Darwin: an intimate of evolution. W.W. Norton and Co., New York, USA. terranes of the Chortis block based on integration of regional aeromagnetic and geologic data. Pp. 65 in: P. Mann (ed.), Geologic and tectonic development of the Caribbean plate in northern Central America. Geological Society of America Special Paper 428. C. LaFemina. 2009. Rough crust subduction, forearc kinematics, and Quaternary uplift rates, Costa Rica segment of the Middle American Trench. Geol. Soc. Amer. Bull. 121: 992-1012. Simkin, T. & R. S. Fisk. 1985. eruption and its effects Smithsonian Institution Press, mineralsciences.si.edu/tdpmap/. 2007. Mexico City, Mexico. Suess, E. 1883-1909. Das Antlitz der Erde http://www.tropicos.org/ accessed 25 January to 8 March, 2009. Trusty, J. L. & M. A. Blanco. 2005. Las orqudeas de la Isla del Coco. Epidendrum 27: 15-18. Fourth Ser., 57: 247-355. An atlas of orchid pollination: America, Africa, Asia, and Australia. A.A. Balkema, Rotterdam, The Netherlands. their pollination and evolution Press, Coral Gables, Florida, USA. Wallace, A. R. 1870. Contributions to the theory of natural selection a series of essays. 1st edition. Macmillan and Co., London, UK. Wallace, A. R. 1889. of natural selection, with some of its applications. 1st edition. Macmillan and Co., London, UK. 96 in: A. G. Coates (ed.). Central America: a natural and cultural history. Connecticut, USA. in: A. G. Coates (ed.). Central America: a natural and cultural history. Connecticut, USA. 2007. Molecular phylogenetics of and related genera (Orchidaceae: Cymbidieae) based on combined molecular data sets. Amer. J. Bot. 94: 18601889. World Checklist of Monocotyledons. 2009. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the Internet; http://www.kew.org/wcsp/monocots/ accessed 25 January to 8 March, 2009. Yeats, R. B., K. Sieh & C. R. Allen. 1997. In The geology Orchid species of Peru.
LANKESTERIANA 11(3): 293. 2011.THE ROLE OF COMMON ORCHIDS IN APPRECIATING THE COMPLEXITY OF BIODIVERSITY CONSERVATION MARILYN H. S. LIGHT & MICHAEL MACCONAILL Corresponding author: email@example.com ABSTRACT. populations of two common terrestrial orchids that grow in close proximity in Gatineau Park, Qubec, Canada. pubescens and of the introduced Epipactis helleborine presented us with an opportunity to conduct in situ experimentation, and that the distribution, patchiness, and persistence of E. helleborine and that seeds of this orchid in trampled soil may germinate better than those in undisturbed soil but that location changes in the soil fungal assemblage upon which the nematodes feed. While mature plants of pubescens not become aware of such changes for some time. RESU M EN. Para conse ser totalmente errneo si no se toman en cuenta aspectos crticos de la historia natural o las relaciones con otros organismos. Por ms de dos dcadas hemos monitoreado las poblaciones de dos orqudeas terrestres y tambin que las semillas de las orqudeas germinan mejor en tierra apisonada que en aquella que no daremos cuenta de estos cambios KEY WORDS : pubescens, Epipactis helleborine
orchid populations in Gatineau Park, Qubec, Canada, one of our concerns was the potential impact of our common orchids, pubescens (Willd.) O.W.Knight and the introduced European orchid, Epipactis helleborine only were there large populations consisting of hundreds to conduct in situ experimentation which might not habitat, we followed a set path within sites and limited how we could assess our physical impact on things we could not see or predict and what measures could be undertaken to mitigate further monitoring impact on the orchid ecosystem. In 2006, we began a study to Cypripedium pubescens was selected for this study because it was shallow-rooted and therefore potentially in pre-set locations for 10 days during the orchid Trampling spots were located where one might stand Soil nematodes were selected as a bioindicator because indicator of soil health and can also be easily extracted study, with follow-up measurements during 2007, soil and perturbed the nematode community. These after 16 months (Light & MacConaill, 2007, 2008). We needed to learn how long the trampling effect of be repeated in different habitats, and if trampling decided to use the seeds of E. helleborine instead of the Cypripedium impact on seed germination because we already had conducted a preliminary study of in situ germination of E. helleborine. Materials and methods Effects of visitor trampling on a terrestrial orchid habitat In 2008, the earlier trampling experiment (Light & MacConaill, 2007) was repeated in three nearby yet different sites in Gatineau Park, Qubec, Canada, that had the following characteristics. Site A was a long-term study site located in open forest (Light 5.8 .0: n=15); dominant trees, Acer saccharum Marshall and Quercus rubra L.; orchid, Cypripedium pubescens. Test and control plots 10 years). Site S was an isolated forest clearing located 6.1 6.8: n=15); dominant tree, Acer saccharum; orchid, pubescens A featuring a centrally located game trail: humus soil dominant trees, Quercus rubra, Pinus strobus L., and Acer saccharum. small population of (Torr.) Lindl. We established experimental plots along either side orchids. Experimental trampling consisted of a 5-minute pubescens in May. Soil compaction and temperature were assessed daily for a total of 30 days before, during, and after the experimental trampling period, and monthly thereafter until September. A pocket penetrometer (Cole-Palmer) was used to measure resistance to penetration of the soil by a standard cylinder. The nematode community LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.294 LANKESTERIANA
composition of all experimental plots and the described (Light & MacConaill, 2007), with paired soil cores (5.5 cm dia x 3 cm deep) being taken from a randomly selected footprint in each sample plot in footprint surface was still present for the compaction measurements in those plots. Distribution and association of Epipactis helleborine with large treesEpipactis helleborine is locally knowledge of its biology and distribution in our longterm study Site 1. It was in this site that we noticed a clear association with a large tree, 44 cm diameter at breast height (dbh), rather than with any of the 31 much smaller trees and saplings (1 to 11.5 cm dbh) that shared the site (Light & MacConaill, 2006a). The 15 largest trees in an area of open forest adjacent to this 2005 (Fig. 1). These trees were 15 to 20 m tall and ranged in diameter (dbh) from 17 to 75 cm with a saplings in this area were mainly Ostrya virginiana (Mill.) K.Koch. All E. helleborine plants emerging during 2006 within a 3 m radius from the trunk of each of the 15 large trees (total area approx. 445 m2) were counted and mapped. For the purpose of comparison, outside those circles (Fig. 1). All orchids emerging in this area were counted and mapped. Trampling during possible on exposed rocks and by keeping to an established assessment trail. Plant densities in selected areas were compared by calculating the X2 statistic (Sokal & Rohlf, 1981) for the differences between expected from the null hypothesis of a uniform Impact of trampling on germination of Epipactis helleborine We used our knowledge of E. helleborine where we expected to obtain the most useful results: were likely present, within 5 m of a large tree. Mature the experiment. Seeds were dried and stored at room before preparation and placement of seed packets mounts. Approximately 150 to 200 seeds were placed in each packet and the mounts stapled at three points. A small hole was punched through one corner of a fastened. All packets could thus be tethered to a central packet-pairs per plot (24 seed packets in all) were buried within 5 m of a large tree, Carya cordiformis (Wangenh.) K.Koch, that had died after a 1998 storm had broken LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. LIGHT & MACCONAILL 295 FIGURE 1. E. helleborine found in 2006, and seed packet burials (1 to 4) within 5 m of a dead Carya cordiformis (C2) and its fallen treetop (solid line). Trees are circumscribed by 3 m radius circles.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.296 LANKESTERIANAthe treetop (Fig. 1 C2). The fallen treetop log formed the upper boundary of packet burials (Fig. 1). Pairs of tethered seed packets were buried in groups of 3 with peg. Two groups were sited 1 m apart and about 50 cm from the log. The two other groups were sited about 1 m apart and away from the others. The packet groups were located between the two clusters of E. helleborine soil disturbance during packet placement, a stout knife was used to cut slots in the soil into which packets were each group of 3 packet-pairs, two pairs were randomly chosen for trampling at a rate of 30 footsteps per packet (16 packets trampled in all). Controls were untrampled. Soil resistance to penetration was assessed before and after packet placement and two years later just before were washed, opened and scored for germination and not split; Stage 2 swollen embryotesta split; Stage root(s). Results Follow up of the 2006 studyResidual soil compaction still persisted 28 months after the initial 2006 study (Fig. 2). Soil compaction measurements still tracked with soil temperature (Fig. 2), with resistance to penetration increasing and decreasing concomitantly with soil temperature. Both trampled and untrampled sampling of trampled plots in the third year. There measure residual compaction. Effects of trampling in different habitatsThe results were essentially the same at each of the 3 sites. Soil compaction increased with each trampling episode during the 10day experimental period and remained for the four months after experimental trampling until on two occasions induced a brief soil softening before Soil nematode assemblages, although different in composition at each site, were similarly impacted by trampling. The experimental trampling and trail t=3.23 (P<0.001)]. Distribution and association of E. helleborine with large treesWe found 159 E. helleborine plants within 3 m Twenty-four plants were found outside the 3 m circles but within the control region: 12 of these plants were the 3 m circles and the control region, one of which FIGURE 2. by gray bar. Soil temperature shown by crosses. FIGURE 3.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. LIGHT & MACCONAILL 297 Carya cordiformis C2 and within 1 m of the fallen top of that tree. associated with some of the large trees, although there X102 = 90, P << 0.001) in the distribution. The greatest density (52 plants in 60 Acer saccharumA2 and the two Tilia americana L., the four next densest being locations around Acer A3, Carya cordiformis C1 and the two pairs of close neighbors, Acer A7 with the Quercus rubra and Carya C4 with the Ostrya virginiana. In the latter two cases, the orchids were concentrated where the 3 m assessment areas regions and 24 in the remaining outer 98 m2 (X2 = 65, P << 0.001), and with 9 of the 17 remaining plants associated with the Quercus being within 4 m of the trunk of the neighboring Acer (Fig.1). Impact of trampling on germination of E. helleborine Three of four seed packet groupings showed a germination (Fig. 4). With these groupings, germination greater with trampled packets than with controls. While control and trampled plots had been similarly loosened when packets of seeds were buried, only the more compacted after two years showed enhanced grouping, located within 2 m of the other groups and close to the felled top of the dead Carya cordiformis, being within 4 m of the trunk of that tree, control and trampled seed packets showed equally high (94%) could not separate the results of control and trampled Discussion The use of common plants and animals as models toward is not surprising. Many of these model organisms were Koormeet, 2002). The nematode, Caenorhabditis elegans Maupas, the mustard, Arabidopsis thaliana Antirrhinum majus mimetic orchids, which employ deception to attract mimicry (Schlueter & Schiestl, 2008). species at risk yet often without an understanding of to shared challenges including disturbance and to Gaston and Fuller (2007). There may be more of a single species and that species may be widespread, but there are not that many species that are so common or Backhouse, 2007). FIGURE 4. lopment by stages in packets of E. helleborine seeds buried at 5 cm depth for 2 years, with and without 30 footsteps immediately after burial.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.298 LANKESTERIANA is not always possible, especially in urban parks attraction. In the Mingan Archipelago National Park a population of Cypripedium passerinum Richardson. colonies were below the minimum plant numbers Cantin, 1998). Because the impact of monitoring on appropriate to monitor the other four colonies. Perhaps the knowledge of trampling impact on the habitat will assist park managers with a similar challenge. Until anthropogenic disturbance on the establishment and nor can we begin to address the challenge of ecosystem Long-term study of common temperate terrestrial orchids in Gatineau Park, Qubec, Canada, has been and germinable seed production of Cypripedium pubescens (Light & MacConaill, would impact the terrestrial ecosystem which could ultimately affect the orchids (Light & MacConaill, 2007, 2008). It is helpful that both E. helleborine and can and do grow in close proximity in Gatineau Park where weather and natural disturbances such as the catastrophic ice storm of January 1998 were shared phenomena yet elicited different responses by these orchids (Light & MacConaill, 2006b). Where a single pubescens growing within the E. helleborine the next 3 years), mature plants of E. helleborine were not similarly affected (Light & MacConaill, 2002). Indeed, the earlier natural death of a large tree in the same part of Site 1 and subsequent drought was suggested as a reason for the decline of plant numbers near that tree, whereas the presence of another healthy large tree about 10 m away was suggested as the reason why some E. helleborine 3 m of that tree to re-emerge after 17 years (Light & MacConaill, 2006a,b). The association of emergent E. helleborine in 2006 with certain species of large trees is therefore not surprising and underscores the importance of such large trees to this common orchid. The exceptional concentration of orchids between tree pairs and with the 3-tree grouping warrants further orchid in North America, seeds do not germinate experience with experimental in situ seed germination (unpublished data) guided the placement of seed packets in the present study. Others working with rare or uncommon species of Epipactis Cephalanthera useful. Our experiment on the impact of trampling on buried seeds of E. helleborine trampling affected germination and initial seedling and trampled plots were similarly loosened when packets of seeds were buried, only the trampled plots a fallen treetop and within 4 m of the trunk of that dead tree, control and trampled seed packets showed equally high germination and enhanced seedling (Bidartondo & Read, 2008). Voss (1972) suggested that E. helleborine Wittig and Wittig (2008) wondered why E. helleborine, compared to so many other terrestrial species, has been Europe. Perhaps soil compaction alters habitat to its across North America since it was introduced in the orchid prefers disturbed ground beside trails may not be ill founded.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. LIGHT & MACCONAILL 299especially so in research trails where this effect has soil (burying seed packets or the insertion of probes on the nematode community or on the germination of E. helleborine as did disturbance by compaction. The data indicate that trampling somehow disrupts the nematodefungus interaction in the compacted community, where orchid seed germination outcomes are determined. This is especially interesting in the case of E. helleborine et al., 2004). Trampling could somehow be modifying et al., 2001; Bidartondo & Read, 2008). Certain fungi in trampled soil could become less palatable to nematodes or less accessible as a food source, which could happen if the fungal cell walls became thickened and less penetrable by nematode feeding stylets. Additionally, nematodes might not reproduce in trampled areas or they could migrate elsewhere if food choices became limited. It is also possible that the soil bacterial community is changed because of trampling and this impacts differentially upon the fungal assemblage or the nematode community, but this has not been (Light & MacConaill, 2007). There is a growing body of concern expressed by common species (Jiguet & Juilliard, 2006; Whiteley et al., 2006; Gaston & Fuller, 2007). There is a real need for understanding of both rare and common species support what are now common species. Gaston and should be heeded by those of us tasked with orchid ACKNOWLEDGMENTS. The authors thank the National Capital Commission for permission to do research in Gatineau Park. LITERA TURE CITED Molec. Ecol. 17: 3707. Brussaard, L., T. W. Kuyper, & R. G. M. de Goede. 2001. fungi and plants: functional composition of species and plant performance Pl. Soil 232: 155. Australian Region Orchid Specialist Group. Third Gaston, K. J. & R. A. Fuller. 2007. Commonness, population 23: 14. in: Press, Florida, USA. Jiguet, F. & R. Julliard. 2006. Inferences from common Biod. Cons. 15: 799. Cypripedium pubescens. Pp. 85 in: P. Kindlmann, orchid populations. Backhuys Publishers, Leiden, The Netherlands. population of pubescens (Willd.) Knight in Gatineau Park, Qubec, 14: 17. disappearance of a weedy orchid. Fol. Geobot. 41: 77. 26: 174. 7: 294. Orchids 77: 128. deoeuf-de-passereau (Cypripedium passerinum) au
Saint-Laurent Vision 2000. their use as indicators. J. Nemat. 33: 161. Schlueter, P. M. & F. P. Schiestl, 2008. Molecular 13: 228. Selosse, M.-A., A. Faccio, G. Scappaticci, & P. Bonfante. 2004. Chlorophyllous and achlorophyllous specimens of Epipactis microphylla (Neottieae, Orchidaceae) Sokal, R. R. & F. J. Rohlf. 1981. Biometry. 2nd edition. W. the history of Arabidopsis Gen. 3: 883. Voss, E. G. 1972. Part I. Gymnosperms and monocots. Cranbrook Institute of Science and Michigan, USA. Whiteley, A. R., P. Spruell, & F. W. Allendorf. 2006. Can 15: 2767. Wittig, R. & M. Wittig. 2007. Epipactis helleborine (L.) Central Europe. Feddes Repert. 118: 46.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.300 LANKESTERIANA
LANKESTERIANA 11(3): 301. 2011.BOTANIC GARDENS, EDUCATION, AND ORCHID CONSERVATION STRATEGIES: THE NEED FOR A COORDINATED APPROACH THOMAS J. MIRENDA Smithsonian Institution, P. O. Box 37012, Greenhouse Nursery Operations, MirendaT@si.edu ABSTRACT how to tackle this worldwide problem. The news has been dire with the prospects of global warming, habitat and technology, and holdings of botanic gardens and responsible commercial growers. It is such horticultural work that has and Wollemia as well as orchid species such as Paphiopedilum vietnamense Epidendrum ilense and Angraecum longicalcar from complete extinction. Botanic gardens, while collections; 2) 50% has been made toward these admirable goals without a unifying entity to compile the progress, information, and successes? With so much effort going on in so many separate places, such an entity is sorely needed. In RESUMEN. diferentes opiniones sobre la manera para atacar este problema global. Las noticias han sido desalentadoras de los Jardines de Lankester, Fundacin EcoMinga, Ecuagenera) y
from disparate disciplines together to work on strategies is increasingly clear that the present generation of in their power to change course before mass extinction occurs (W. F. Laurance, unpublished). In the animal adapted that could help the botanical community create awareness of some of the successes and failures own efforts to protect imperiled plants. The orchid community, by nature of the passion gneros que no corresponden a las orqudeas, tales como Torreya, Wollemia, y especies de orqudeas como Paphiopedilum vietnamense Epidendrum ilense, y Angraecum longicalcar de la extincin. Los jardines colecciones en tantos lugares separados, es indudable que se necesita urgentemente este tipo de entidad. Constituye la meta KEY WORDS LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.302 LANKESTERIANA
history of rampant wild collection, habitat destruction, Ecominga Foundation (http://ecominga.net/) supported numerous projects around the world (Important Plant Areas), that are natural hotspots for same goals. As admirable as these efforts are, more coordination, collaboration, and focus are necessary to efforts are necessarily dependent on the success of reforestation and habitat reconstruction projects. return extirpated orchids to their original wild habitats, then such efforts are futile when those habitats no longer exist. Without a suitable ecosystem to support it, including the entire milieu of plants, insects, and Costa Rica. Though not as ideal for habitat as primary forest, such forests can still contain up to 80% of the tree species found in primary forest. If left alone or supplemented with absent species, these forests could still be excellent habitats for reintroduction of orchids. between primary forests and agricultural monocultures. enjoyment, and education. This contribution should not be underestimated as it is likely the only opportunity for present-day leaders in the botanical community got their and highschool textbooks should pay off in greater Such garden collections, if properly documented and kept disease-free, are also repositories for for genetic material for breeding programs. www.bgci.org) and the American Public Garden Association (APGA; http://www.publicgardens.org/) details such as the need for translations into English are often keeping thousands of species from being redwere adopted from the Global Strategy for Plant 1. 90% of all threatened orchid species should be secure in collections. While progress admirable goal, it seems certain that with so many endemic species in countries such as Ecuador that placed close to 90% of threatened orchid species in situations. It is incumbent on institutions in and sent to appropriate botanic gardens to ensure LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. MIRENDA 303
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.304 LANKESTERIANA programs are considerably more complicated years ago. The need to restore forest (and other) them requires the participation of a large number of biologists to understand fully the implications orchids in Madagascar, the Ecominga foundation and Panama. 3. No orchids should be threatened by unsustainable wild orchids has been curtailed by the enforcement International Trade in Endangered Species persists in the orchid world. The most recent the habitats of Paphiopedilum vietnamense (O.Gruss & Perner) and horticulturally important plants were plundered mercilessly by commercial orchid companies and P. vietnamense, the local authorities were unable to control or enforce laws against wild collection, In addition, smuggled plants acquired by CITES rescue centers were propagated in the hope that reduce wild-collection pressure. Although this effort was successful, it also created a smokescreen for many illegally collected plants to enter the Phragmipedium in the protection and propagation effort for this the engagement of local stakeholders will usually result in better stewardship of endangered species (including orchids) by 2010. Although reaching admirable progress has been made on this resolution. With the worldwide consciousness our natural systems, it seems likely that the next generation of students will be keenly aware of the importance of biological (including botanical) (North American Plant Collections Consortium) promise a future database where germplasm from a and used for breeding purposes. Accreditation by the possessing endangered species in their collection participating institutions are required to follow their endangered holdings. These are associated with databases that record husbandry and collection data the plant resources in botanic gardens and other orchid species collections. Much work has been done in the U.K. and U.S. Projects such as the OSSSU (Orchid Seed Stores for Sustaniable Use) and the Orchid Seedbank Project Space requirements would certainly limit the quantity
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. MIRENDA 305 projects. Other lessons to be learned from the animal world include an analysis of their prowess at outreach, fund-raising, and public engagement. The Internet tremendous interest in orchids that clearly exists when culture and discussion are considered. The website amphibianark.org) is a prime example of how to use the Internet as a fund-raising tool, building the awareness of the need for public engagement and reaching out to many constituencies, especially children. Another understood and promulgated among more people. And, pollination strategies, and charismatic young people should be producing and promoting programming that will engage the public, raise their consciousness, and Perhaps most importantly, it has become apparent success. The case history of the Takhi (the Mongolian wild horse) is a prime example of a reintroduction fully taking into consideration its foraging needs the dominant male and his females, the rest of the expertise to restore the habitat of the Takhi prior to It will always be necessary for animal and plant the same way that plants and animals, on the most for grants) together that address the interconnected LITERA TURE CITED 23: 1406-1417. Electronic sources: http://nationalzoo.si.edu/ConservationAndScience/CRC/ np accessed 10/ 2008 http://www.orchidconservation.org/OSG/ np accessed 10/2008 http://www.orchidconservationalliance.org/ np accessed 10/2008 http://www.publicgardens.org/ np accessed 10/2008 http://www.cbd.int/gspc/ np accessed 10/2008 http://ecominga.net/ np accessed 10/2008
part. For part of its nomenclatural history it has been known as subtribe Sobraliinae (although placed in subtribe Sobraliinae in tribe Arethuseae based on LANKESTERIANA 11(3): 307. 2011.PRELIMINARY MOLECULAR PHYLOGENETICS OF SOBRALIA AND RELATIVES (ORCHIDACEAE: SOBRALIEAE) KURT M. NEUBIG 1,2,5 W. MARK WHITTEN 2 MARIO A. BLANCO 1,2,3 LORENA ENDARA 1,2, NORRIS H.WILLIAMS 2 & SAMANTHA KOEHLER 412 Florida 32611-7800, U.S.A.34 5 ABSTRACT. As currently circumscribed, the tribe includes four genera: Elleanthus, Epilyna, Sertifera and Sobralia. Most Elleanthus, Epilyna and Sertifera but not Sobralia Sobralia. within the tribe. With few exceptions, species of Sobralia predominantly offer no reward and are pollinated by bees. Elleanthus and Sertifera rewards. Nothing is known of pollination in Epilyna RESUMEN. Elleanthus, Epilyna, Sertifera y Sobralia. Las plantas de stos cuatro gneros generalmente producen tallos largos como caas, pero Elleanthus, Epilyna y Sertifera pero no de Sobralia. Sobralia Sobralia. Sobralia no Elleanthus y Sertifera Epilyna KEY WORDS : Orchidaceae, Sobralieae, Sobralia, phylogenetics
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.308 LANKESTERIANAsymplesiomorphies such as presence of corms, plicate aberrant genera such as Arpophyllum and Xerorchis). placement of Sobralieae has been associated with other basal members of subfamily Epidendroideae based on plesiomorphic subfamilial characters. More recent and demonstrated that Sobralieae are basal members of the subfamily Epidendroideae, closely related to genera such as Tropidia (Cameron et al., 1999; Cameron, 2002, 2004). Because this group is not closely related to other taxa in tribes Epidendreae and Arethuseae, the tribe (see Pridgeon et al., 2005). Tribe Sobralieae consists of only four genera of unequal species richness. Two genera, Elleanthus C.Presl. and Sobralia about 100 species, whereas the other two genera, Epilyna Schltr. and Sertifera Lindl. & Rchb.f., each consist of less than 10 species. The tribe as a whole is widely distributed in tropical America. Sertifera is Andes. Epilyna is found in southern Central America and northern South America. Elleanthus is distributed throughout tropical America, and Sobralia is similar in distribution except for notable absence in the West Indies. identifying species or groups within Sobralieae, there (Fig. 1). Sobralia (Elleanthus, Epilyna, Sertifera has been shown to result in the circumscription of polyphyletic groups based on homoplasious character et al., 1998). Because there has been such a poor understanding of generic circumscription in Sobralieae and no robustly taxonsampled phylogenetic analysis of the tribe, we addressed phylogenetic relationships within the tribe. for reciprocal monophyly in these genera because the polarity of such a character would make one state symplesiomorphic. Therefore, the purpose of this study tribe Sobralieae. Materials and methods Specimens were obtained from Sampling of Elleanthus, Epilyna, Sertifera, and Sobralia included 42 species. Outgroups included three other genera of basal Epidendroid tribes Neottieae (Palmorchis), Arethuseae (Bletilla), and Tropidieae (Tropidia). Outgroups were chosen based on phylogenetic placement of Sobralia and Elleanthus et al., 1999; Cameron, 2002; Chase et al., 2003; Cameron, 2004). All freshly were ground in 1 mL of CTAB 2X buffer and either performed using a Biometra Tgradient or an Eppendorf Mastercycler EP Gradient S thermocycler and Sigma units Taq For the plastid regions the following reaction Taq parameters 94 C, 2 min; 15X (94 C, 1 min; 76 C, 1 min, reducing 1 C per cycle; 72 C, 1 min); 21X (94 C, 1 min; 59 C, 1 min; 72 C, 1 min); 72 C, 3 min with the
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. NEUBIG et al. Preliminary phylogenetics of Sobralia309primers 17SE (ACG AAT TCA TGG TCC GGT GAA GTG TTC G) and 26SE (TAG AAT TCC CCG GTT CGC TCG CCG TTA C) from Sun et al. (1994). trnSGCU-trnGUCC the parameters 94 C, 3 min; 33X (94 C, 30 sec; 50 C, 30 sec; 72 C, 2 min); 72 C, 3 min, with the primers trnSGCU (AGA TAG GGA TTC GAA CCC TCG GT) UUC (GTA GCG GGA ATC GAA CCC GCA TC) from Shaw et al. (2005). ycf1 We sequenced a ca. 1500 base-pair (bp) portion et al., 2009). This region was parameters 94 C, 3 min; 8X (94 C, 30 sec; 60-51 C, 1 min; 72 C, 3 min); 30X (94 C, 30 sec; 50 C, 1 min; 72 C, 3 min); 72 C, 3 min, with primers 3720F (TAC GTA TGT AAT GAA CGA ATG G) and 5500R (GCT GTT ATT GGC ATC AAA CCA ATA GCG). Additional internal primers intF (GAT CTG GAC CAA TGC ACA TAT T) and intR (TTT GAT TGG GAT GAT CCA AGG) were also required for sequencing. PCR products were cleaned with Microclean (The Gel Company, San Francisco, CA, USA) following FIGURE Sobralia, such as in A) S. citrea, B) S. callosa, C) S. crocea S. luerorum. Various members of Sobralia sect. Sobralia include E) S. ciliata F) S. portillae G) S. mandonii S. caloglossa (not sampled in this study, but unpublished data place this species in a clade with S. mandonii and S. dichotoma). Most members of the genus Elleanthus E. caravata E. lancifolius K) Species of the genus Sertifera
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.310 LANKESTERIANA TABLE
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. NEUBIG et al. Preliminary phylogenetics of Sobralia311 PCR products were then cycle-sequenced using the parameters 96 C, 10 sec; 25X (96 C, 10 sec; 50 C, sequencing products were directly sequenced on an ABI 377, 3100 or 3130 automated sequencer according Foster City, CA, USA). Electropherograms were edited and assembled using Sequencher 4.9 (GeneCodes, Ann Arbor, MI, USA). All sequences were deposited in GenBank (Table 1). Data analysis Sequence data were manually aligned data were excluded from analyses. Indels (insertions/ deletions) were not coded as characters. Analyses were performed using PAUP*4.0b10 (Swofford, 1999). Fitch parsimony (unordered characters with equal weights; Fitch, 1971) analyses used a heuristic search strategy consisted of branch swapping by addition replicates holding 5 trees at each step, and were assessed using the bootstrap (Felsenstein, 1985). Bootstrap percentages under parsimony were estimated with 1000 bootstrap replicates, using TBR swapping for 50 randomaddition replicates per bootstrap replicate. For maximum likelihood (ML), Modeltest (Posada & Crandall, 1998) was used to determine the appropriate model for analysis using all combined data under the Akaike Information Criterion. ML analyses were set. Bootstrap percentages under ML were estimated with 100 bootstrap replicates, using TBR swapping for one randomaddition replicate per bootstrap replicate. All analyses were performed for data sets including congruence was tested using the partition homogeneity
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.312 LANKESTERIANA replicates and TBR branch-swapping. Probability Results The aligned length of the ITS data set was 892 bp. Fitch parsimony analysis of the ITS region found 100 equally parsimonious trees of 798 steps (consistency index (CI) = 0.589, retention index (RI) = 0.753). The aligned length of the combined plastid data set (trnS-G and ycf1) data set was 2919 bp. Of these, 250 analysis of the combined plastid data set found 100 equally parsimonious trees of 1112 steps (CI = 0.772, RI = 0.794). The aligned length of the combined (three trnSG, and ycf1) was 3811 bp. Of these, 472 were potentially parsimony1926 steps (CI = 0.690, RI = 0.767). Maximum likelihood analysis of ITS only (not presented), plastid data only (not presented), and all three regions (-lnL = 16599.46) yielded trees similar in topology to parsimony. Bootstrap support for all nodes was similar to that from parsimony. The only exception Sobralia ciliata in plastid Partition homogeneity tests showed mixed results for congruence among the different partitions of these data. The test comparing ITS and the combined plastid FIGURE ycf1 and trnS-G than 50%.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. NEUBIG et al. Preliminary phylogenetics of Sobralia313 incongruence (ITS/trnS-G P=0.10; ITS/ycf1 P=0.13; ycf1/trnS-G percentages between the different data sets (Fig. 2) indicates that there are only a few examples of strong incongruence. For example, Sobralia ciliata is sister Sobralia according to ITS but sister to the rest of the tribe in the plastid data set. positions of S. dorbignyana, S. portillae, S. mandonii, S. dichotoma, and Sertifera colombiana. All data were combined because the partition homogeneity (Graham et al et al., 2001) and because With limited outgroup taxon sampling, relationships among the basal Epidendroideae tribes Neottieae (Palmorchis), Tropidieae (Tropidia), Arethuseae (Bletilla tribe Sobralieae is monophyletic in all data sets. Within Sobralieae, there are many consistent Sobralia (see Fig. 3, 4), Elleanthus, and Epilyna are all consistently monophyletic. Because only one sample of Sertifera was used in this study, monophyly of the genus could not be determined. Inconsistent features of phylogenetic topology are centered on Sobralia species within section Sobralia: S. dichotoma, S. ciliata, S. dorbignyana, S. mandonii, and S. portillae. among different data sets. Discussion Morphological characters supporting the monophyly of Sobralieae include an elongate cane-like stem Within Sobralieae, Elleanthus and Epilyna are both monophyletic, but Sobralia is polyphyletic. We sought morphological features that might distinguish the in Sobralia. These features are discussed below. are formed at the apex of a shoot and axillary the shoot terminus. The distinction between these two positions can be blurred in some plant groups, but in Sobralieae, the difference is usually clear (see Fig. 1, in a few species (e.g., Sobralia dorbygniana), both rachis) may be highly condensed (capitate in some species of Elleanthus) or elongate, branched or unbranched, erect or (less commonly) nodding, and In a few species of Elleanthus In Sobralieae, all of these inflorescence These differences are presented in the simplified illustrations of Figure 4. As delimited in Figure 3, Sobralia main types of inflorescence morphology. Both types are terminal, but in species such as S. rosea and S. luerorum ( S. sect. Racemosae) the floral displays are strongly distichous and the rachis is fractiflex Sobralia liliastrum also has this inflorescence morphology, and when combined with S. rosea and S. luerorum, this assemblage is paraphyletic. In the remainder of Sobralia condensed, such that the internodes of the rachis are extremely short (often 1-2 mm). The resulting large bracts. This condensed inflorescence is present in many Sobralia with ephemeral flowers. In the combined analysis (Fig. 3, 4), Sobralia ciliata S. dichotoma and S. mandonii are sister to the remainder of the tribe. S. sect. Sobralia. In addition to the genus Sertifera, these
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.314 LANKESTERIANAS. sect. Sobralia ( S. dorbignyana and S. portillae) Epilyna and Elleanthus. Elleanthus the whole tribe. Elleanthus distichous or spirally arranged, capitate to loosely racemose, and can be oriented downwards, upwards or clades of Sobralieae demonstrate the plesiomorphic symplesiomorphic grade across both major clades is represented by some taxa of S. sect. Sobralia and Sertifera. The result is that there has been independent large clades in Sobralieae. Flower size of Sobralieae. Species of Elleanthus, Epilyna, and Sertifera Sobralia likely a consequence of shifts in pollination mode. Sobralia are mostly pollinated by large bees (e.g. Eulaema FIGURE regions (ITS, trnS-G, and ycf1 ).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. NEUBIG et al. Preliminary phylogenetics of Sobralia315 FIGURE trnS-G, and ycf1 ), to demonstrate n.b., Elleanthus are always terminal and consisting of a single axis as indicated by the illustration). Elleanthus and Sertifera are usually pollinated by Epilyna Elleanthus, are unknown. Variation of different pollinators and associated in some systems (Thomson and Wilson, 2008). for shifts in pollination syndrome. Often, species relationships due to homoplasy in pollinationrelated within Sobralia. Sobralia callosa has been segregated as Lindsayella Ames & C.Schweinf. because of its
of most species of Sobralia of Lindsayella in Sobralia connote reciprocal monophyly. generic concepts should not be based primarily diagnostic, and we recommend that future generic recircumscriptions be based on the combination of these apomorphic characters in conjunction with molecular data. Unfortunately, the type species of Sobralia is S. dichotoma (designated by Angely in Fl. Analtica So Paulo 6: 1268. 1973). This species Sobralia paper. As a result of this quirk of history and because of the polyphyly of Sobralia, there are problematic be the subject of future research. ACKNOWLEDGMENTS. We thank Jardn Botnico Lankester specimens and tissue. We are grateful to the Portilla family of Ecuagenera Ltd. in Gualaceo, Ecuador, and to for generous access to their collections. Some specimens in the Life Sciences funded by grants from the U.S. National Interdisciplinary Center for Biotechnology Research at primarily by Kent Perkins at the FLAS herbarium in the were funded by the Lewis and Varina Vaughn Fellowship in Orchid Conference Fellowship to K. Neubig, and the U.S. LITERATURE CITED Cameron, K. M., M. W. Chase, W. M. Whitten, P. J. Kores, D. C. Jarrell, V. A. Albert, T. Yukawa, H. G. Hills, D. H. Goldman. 1999. A phylogenetic analysis of the Orchidaceae: evidence from rbcL nucleotide sequences. Amer. J. Bot. 86: 208-224. Cameron, K. M. 2002. Molecular systematics of Orchidaceae: a literature review and an example using in: H. Nair (ed.). Proceedings of the 17th World Orchid Conference. Natural History Publications (Borneo) Sdn. Bhd., Sabah, Malaysia. Cameron, K. M. 2004. Utility of plastid psaB gene sequences for investigating intrafamilial relationships within Orchidaceae. Molec. Phylogen. Evol. 31: 11571180. Chase, M. W., J. V. Freudenstein, K. M. Cameron & R. L. Barrett. 2003. DNA data and Orchidaceae systematics: in: K. W. Dixon, S. P. Kell, R. L. Barrett & P. J. Cribb (eds.). Orchid conservation. Natural History Publications, Kota Kinabalu, Malaysia. Chase, M. W. & H. G. Hills. 1991. Silica gel: an ideal studies. Taxon 40: 215-220. Doyle, J. J. & J. L. Doyle. 1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull. 19: 11-15. Dressler, R. L. 1981. The orchids: natural history and Harvard University Press, Cambridge, Massachusetts, USA. Dressler, R. L. 1993. orchid family Dioscorides Press, Portland, Oregon, USA approach using the bootstrap. Evolution 39: 783-791. 20: 406-416. Graham, S. W., J. R. Kohn, B. R. Morton, J. E. Eckenwalder & S. C. H. Barrett. 1998. Phylogenetic congruence and discordance among one morphological and three molecular data sets from Pontederiaceae. Syst. Biol. 47: 545-567. Johnson, L. A. & D. E. Soltis. 1998. Assessing congruence: empirical examples from moleculardata. Pp. 297-348 in: D. E. Soltis, P. S. Soltis & J. J. Doyle (eds.). Molecular systematics of plants II: DNA sequencing. Kluwer Academic Publishers, Boston, Massachusetts, USA. Johnson, S. D., H. P. Linder & K. E. Steiner. 1998. Phylogeny and radiation of pollination systems in Disa (Orchidaceae). Amer. J. Bot. 85: 402-411. Neubig, K. M., W. M.Whitten, B. S. Carlsward, M. A. Blanco, L. Endara, N. H. Williams & M. Moore. 2009. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.316 LANKESTERIANA
Phylogenetic utility of ycf1 in orchids: a plastid gene more variable than matK. Pl. Syst. Evol. 277: 75-84. Posada, D. & K. A. Crandall. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817-818. Pridgeon, A. M., P. J. Cribb, M. W. Chase & F. N. Rasmussen (eds.) 2005. Genera orchidacearum, Vol. 4. Epidendroideae (Part one). Oxford University Press, UK. Rambaut, A. 1996. Se-Al: Sequence alignment editor, v2.0a11. Oxford University, Oxford, UK. Available at website, http://evolve.zoo.ox.ac.uk/, last accessed 8 August 2002. Reeves, G., M. W. Chase, P. Goldblatt, P. Rudall, M. F. Molecular systematics of Iridaceae: evidence from four plastid regions. Amer. J. Bot. 88: 2074. Shaw, J., E. B. Lickey, J. T. Beck, S. B. Farmer, W. Liu, J. Miller, K. C. Siripun, C. T. Winder, E. E. Schilling & R. L. Small. 2005. The tortoise and the hare II: relative utility of 21 noncoding chloroplast DNA sequences for phylogenetic analysis. Amer. J. Bot. 92: 142-166. Sun, Y., D. Z. Skinner, G. H. Liang, & S.H. Hulbert. 1994. Phylogenetic analysis of Sorghum and related taxa using internal transcribed spacers of nuclear ribosomal DNA. Theor. App. Genet. 89: 26-32. Swofford, D. L. 1999. PAUP*: phylogenetic analysis using parsimony (*and other methods), version 4.0b10. Sinauer Associates, Sunderland, Massachusettts, USA. Thomson, J. D. & P. Wilson. 2008. Explaining evolutionary shifts between bee and hummingbird pollination: convergence, divergence, and directionality. Int. J. Pl. Sc.169: 23-38.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. NEUBIG et al. Preliminary phylogenetics of Sobralia317
essentially identical, but their transcription program stage, age, location, etc. This critical feature of of epigenetic signals that are not encoded in the access to underlying genetic information to modulate the identity of cells. imprinted gene expression (Steimer et al. 2004, (epialleles) also control the organismic response et al. 2005) LANKESTERIANA 11(3): 319. 2011.EPIGENETIC INFORMATION UNEXPLORED SOURCE OF NATURAL VARIATION OVIDIU PAUN & MARK W. CHASE ABSTRACT. Dactylorhiza traunsteineri D. majalis, and D. ebudensis D. fuchsii and D. incarnata, but still appear genetically uniform may be the result of altered epigenetic controls of gene expression without any change in the underlying genetic material. RESU M EN. genticamente similares. Por ejemplo, sabemos que Dactylorhiza traunsteineri, D. majalis, y D. ebudensis D. fuchsii y D. incarnata, pero stas tienen ecologa y distribucin detectar patrones que indican que algunas de estas diferencias se deben a efectos epigenticos, los cuales ecolgicamente diferentes, pero que son al parecer genticamente uniformes, podran ser el resultado de la cambio en el material gentico. KEY WORDS : orchids, epigeny
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.320 LANKESTERIANAacross generations through mechanisms that are not g cell differentiation within an organism, epigenetics This may be particularly true for plants, where redundant copies of many genes are spread across the genomes as a result of ancient whole-genome et al. may be particularly important for immobile organisms, such as plants, that generally cannot respond within examples of spontaneous heritable epialleles that demonstrably affect key phenotypic characters, such lineages in higher plants. our understanding of the natural mechanisms Bossdorf et al. such as the extent and structure of epigenetic dormant mobile elements and silence redundant cellular processes (see Paun et al. 2007, for a the stability of these epigenetic changes and their that epigenetic, rather than genetic, differentiation The case of Dactylorhiza allopolyploids Dactylorhiza, a genus of temperate terrestrial Dactylorhiza allotetraploid (2n and geographically widespread parental lineages: the diploid (2n = 40) marsh-orchid, D. incarnata (L.) So s.l., and the spotted orchid, D. maculata (L.) So s.l. (including, among others, diploid D. fuchsii et al. 2001, Pillon et al. et al. 2008). As a result, the taxonomy of many dactylorchids is widely considered complicated, causing long-standing ecological requirements and distinct distributions. progenitor pair can exhibit contrasting morphological and ecological properties that are maintained in spite Examples of such cases include European allotetraploids D. majalis & Summerh. s.str., D. traunsteineri (Saut. ex Rchb.) So s.l. (Fig. 1) and D. ebudensis (Wief. endemic, which has most probably a single origin) D. fuchsii (in all cases the maternal parent) and D. incarnata (Pillon et al., 2007, and references therein).
ITS alleles, and in agreement with the patterns of morphology and ecological preference, D. majalis inferred to be the oldest of the three allotetraploids bottlenecks in southern Eurasia. It has a fairly wide ecological tolerance of soil moisture and occurs at present in damp meadows and fens in western and central Europe, the Baltic region, and northern Russia. In contrast, D. traunsteineri is a more heterogeneous and still maintains both parental ITS alleles (Pillon et al., 2007). It probably originated and disjunct distribution in northwestern and central and grows in calcareous fens and marshes. A third allotetraploid, D. ebudensis, is a narrow endemic (at present forming a single population) in northwestern Scotland and may be as young as or younger than D. traunsteineri. The coastal dune habitat occupied by D. ebudensis ness, neutral genetic differentiation between sibling Dactylorhiza allopolyploids, as studied with ITS sequences, AFLPs, and plastid and nuclear et al., 2001; Pillon et al., 2007). By contrast, et al., 2007) resulted in a geographic structure (Paun et al., 2010), grouping the multiple independent origins of each tetraploid taxon (except D. ebudensis discriminate between the allopolyploid species (Paun et al., 2010). the genome, which was studied using methylationseparates the three allopolyploids studied here (Fig. 2), in stark contrast to patterns of genome-wide genetic et al., 2001). The MSAP technique is similar to standard AFLP (Vos et al., 1995) but uses batches (Baurens et al. methylation status at each restriction site. Unexpectedly, methylation status analysis at 332 genome-wide MSAP markers resulted in speciesPillon et al., 2007). Indeed, the migration-induced genetic bottleneck triggered within D. majalis by the profound change of climatic conditions during LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PAUN & CHASE Epigenetic information321 FIGURE 1. The allotetraploid Dactylorhiza traunsteineri at a natural site in Yorkshire, Britain (Photo: O. Paun).
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.322 LANKESTERIANAweak within-species geographic differentiation (Fig. 2). As result of its postglacial formation and present disjunct distribution, the genome-wide methylation patterns of D. traunsteineri are more heterogeneous and correlate more clearly with geography. Although containing information from both coding differences in the three allopolyploid siblings than P < 0.01). Therefore, it seems that habitat preference shapes similar expression patterns in some, but not all, of the independent et al. in press). To test further our hypothesis of adaptation performed scans for outlier epigenetic loci, looking regressions between epigenetic data and ecoet al., 2007); and 2) a Bayesian outlier locus approach as implemented in BayeScan (Foll and Gaggiotti, 2008). The latter approach estimates the posterior probability of each locus being under selection, and it is able to differentiate the type of selection each SAM pinpointed 14 methylation markers as being approaches. For most of the outlier epiloci, their presence-absence patterns correlated with water FIGURE P = 0.001) of methylation status of allotetraploid D. majalis (black symbols), D. traunsteineri (gray symbols), and D. ebudensis line encloses samples from Yorkshire, England, and the dashed line samples from N.W. Scotland.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PAUN & CHASE Epigenetic information323 of drift (Nosil et al., 2009) and/or further selection. Therefore, epigenetic regulatory processes play of the allopolyploid Dactylorhiza species by in epigenetic patterns indicates that such markers rapidly radiating groups when genetic markers may ACKNOWLEDGEMENTS. O.P. was funded by an IntraEuropean Marie Curie Fellowship (EU Comission, MEIFCT-2007-040494). LITERA TURE CITED CCGG methylation in the banana genome. Pl. Molec. Biol. Reporter 21: 339-348. Bossdorf, O., C. L. Richards & M. Pigliucci. 2008. Epigenetics for ecologists. Ecol. Letters 11: 106-115. Foll, M. & O. Gaggiotti. 2008. A genome-scan method to identify selected loci appropriate for both dominant and 180: 977-993. 2005. Epigenetic differences arise during the lifetime 10604-10609. epigenetic control of gene expression. Science 301: 798802. world of small RNAs. Nature 451: 414-416. Dactylorhiza 201: 31-55. Dactylorhiza (Orchidaceae). Amer. J. Bot. 88: 1868-1880. Dactylorhiza incarnata/maculata complex (Orchidaceae) in Molec. Ecol. 17: 5075-5091. inheritance in plants. Nature 447: 418-424. Naturwiss. Ver. Wuppertal Biol. 84: 131-176. Joost, S., A. Bonin, M. W. Bruford, C. Conord, G. Erhardt & P. Taberlet. 2007. A spatial analysis method (SAM) to detect candidate loci for selection: towards a landscape genomics approach to adaptation. Molec. Ecol. 16: 3955-3969. fungi. Science 293: 1070-1074. Molec. Ecol.18: 375-402. L. and Chase, M. W. In press. Stable epigenetic effects and adaptation in allopolyploid orchids (Dactylorhiza: Orchidaceae). Molecular Biology and Evolution doi: and genome doubling. 56: 649-656. Paun, O., J. A. Luna, M. F. Fay, R. M. Bateman & M. in allotetraploid species of Dactylorhiza (Orchidaceae; Orchidinae). Pp. 169-192 in: O. Seberg, G. Petersen, A. of western European polyploid species complexes in Dactylorhiza (Orchidaceae). 56: 1185-1208. Rapp, R. A. and Wendel, J. F. 2005. Epigenetics and plant 168: 81-91. 7: 395402. P. K. Wall & P. S. Soltis. 2009. Polyploidy and angiosperm
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.324 LANKESTERIANA Current Opinion Pl. Biol. 7: 11-19. 23: 4407-4414.
LANKESTERIANA 11(3): 325. 2011.OF GREENISH ENCYCLIA: NATURAL VARIATION, TAXONOMY, CLEISTOGAMY, AND A COMMENT ON DNA BARCODING FRANCO PUPULIN 1-5 & DIEGO BOGARN 1-4123 Marie Selby Botanical Gardens, Sarasota, Florida 34236 USA45 Corresponding author: firstname.lastname@example.org ABSTRACT. Encyclia has been disputed considerably because of the great morphological similarity among many of the taxa, particularly in the complex of species related to E. chloroleuca and E. gravida the natural lineages of the greenish species of Encyclia and species concepts in this group are discussed here E. amanda to synonymy of E. chloroleuca Encyclia referred to E. gravida of E. gravida may simply represent self-pollinating forms belonging to different taxa. RESU M EN. Encyclia para un amplio complejo de especies emparentadas con E. chloroleuca grupo reconocieron solamente cuatro especies (con cinco subespecies). Ambas interpretaciones, sin embargo, se basaron fundamentalmente en el estudio de material escaso, a menudo unas pocas colecciones por cada pas, Encyclia probablemente no descrita) algunas de las cuales se extienden a Suramrica. La taxonoma de los taxones de separar con mtodos taxonmicos tradicionales. Sin embargo, los cdigos de barras por si solos nunca son
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.326 LANKESTERIANA KEY WORDS : Orchidaceae, Laeliinae, Encyclia Central America, taxonomy, cleistogamy The need for rigorously delimited species concepts described organisms increases at an unprecedented rate, our knowledge of many of the old and new taxa is still based on a minimal number of samples. which are in many cases only fragmentarily known geographic ranges. The proliferation of synonyms in species descriptions, which traditionally plagued and between populations on the basis of the limited Flora Neotropica in the 1960s (to date, only the genera Bulbophyllum, Epidendrum, Isochilus, Mormodes, and Vanilla for inclusion) and Flora Mesoamericana in the last where they naturally occur. In the case of the Neotropical orchid genus Encyclia more natural way to circumscribe the genus (e.g., Lindley 1842, 1853, Ames et al. et al. et al. 2003), but the task of describing the Encyclia any scheme of relationships among species. Joseph Encyclia in 1828 based on a Cattleya, the type species for the genus, is indeed a peculiar and poorly known plant with many column, the lateral lobes of the lip almost completely fused with the midlobe, and a nonforcipate basal midpoint of the 19th century, under the authority of John Lindley, the genus was merged under Epidendrum L., and the type species was assigned to Figure 1. Original illustration of from Cur tiss Botanical Magazine 1828, pl. 2851.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PUPULIN & BOGARN Greenish Encyclia327E. subgenus Encyclium Lindl. section Hymenochyla Lindl. (Lindley 1831, Encyclia by Rudolf Schlechter (1914, 1918, 1922, 1923), it was not until 1961 when Robert L. distinctness of the genus and the need to segregate it from Epidendrum narrower circumscription approximating the original FIGURE 2. Labella of Epidendrum, Encyclia and Prosthechea species. A B E. ciliare; C EncycliamooreanaE. chloroleuca ; E -Prosthechea cochleata ; F P. spondiada. Scale bar = 1 cm. FIGURE Epidendrum, Encyclia and Prosthecheaumns. A Encyclia mooreana; B E. chloroleuca; C Prosthechea spondiada P. cochleata Encyclia mooreana; F E. chloroleuca; G Prosthechea spondiada P. cochleata. Scale bar = 1 cm; double bar = 5 mm. FIGURE Epidendrum, Encyclia and Prosthechea species, showA B E. ciliare; C Encyclia mooreana E. chloroleuca ; E Prosthechea spondiada. Scale bar = 1 cm.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.328 LANKESTERIANA bulk of species now assigned to Prosthechea Knowl. Encyclia identity of many of the species in the genus remains 1998, 2001). The taxonomy of greenish Encyclia taxonomy of Encyclia has been widely disputed, largely because of the great morphological similarity among many of the taxa particularly in the large complex of species related to E. chloroleuca and E. gravida (Lindl.) Schltr., which are and the application of old names often based on poorly the natural lineages of the greenish species, so we feel free to discuss this group independently from nomenclatural uses. E. adenocaula & Lex.) Schltr., E. cordigera E. phoenicea (Lindl.) Neumann), species of Encyclia petals, a distinctly trilobed lip with the lateral lobes erect as longitudinal-radiating, sometimes prominent keels, and a basal, forcipate callus; the column is which commonly embrace the isthmus that separate the lateral lobes from the midlobe. The perianth parts are frequently greenish or pale tan, with the Encyclia corresponding to this scheme are broadly distributed in the Neotropics, from Florida, Mexico, and the West Indies, throughout Central America, to Argentina and South America (Table 1). For plants located in Mesoamerica, which we few years, traditional taxonomic approaches to understanding Encyclia many of the described species of Epidendrum and Encyclia from Mesoamerica as a single polymorphic alliance (Ames et al., 1935, 1936) to the recognition According to the different taxonomic treatments, the group of greenish Central American Encyclia includes 4 to 13 species. The use of broad concepts distinguish between closely allied species reached its Epidendrum (including Encyclia) of North and Central America (Ames et al., 1936), they considered most of the Epidendrum oncidioides American species under a few of the oldest names the pronounced polymorphism of many Central American species, induced some botanists to on subtle differences in type specimens that do not hold up in comparison with broader series of records (Withner, 1998, 2001). In both approaches, the impossibility of fully appreciating the range of
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PUPULIN & BOGARN Greenish Encyclia329 TABLE Encyclia Species Distribution Encyclia acuta Encyclia acutifolia (Sw.) Nir Jamaica Encyclia alanjensis Encyclia amanda Encyclia amicta Encyclia angustiloba Encyclia argentinensis Encyclia aspera (Lindl.) Schltr. Colombia, Ecuador, Peru Encyclia asperirachis Garay Colombia Encyclia asperula Encyclia bradfordii Encyclia brenesii Schltr. Costa Rica Encyclia ceratistes Encyclia chloroleuca Encyclia confusa Encyclia davidhuntii Encyclia diurna Encyclia fehlingii (Sauleda) Sauleda & Adams Bahamas Encyclia fucata (Lindl.) Britt. & Millsp. Bahamas, Cuba Encyclia glandulosa Encyclia gonzalensis Encyclia goyazensis Encyclia granitica Encyclia guentheriana Encyclia guyanensis Encyclia hunteriana Schltr. Panama Encyclia inaguensis Nash ex Britt. & Millsp. Bahamas, Turks and Caicos Islands Encyclia leucantha Encyclia lineariloba Withner Mexico. Guatemala, Nicaragua Encyclia linearifolioides Encyclia longifolia Encyclia maderoi Schltr. Colombia Encyclia maravalensis Withner Trinidad Encyclia monticola Encyclia mooreana Encyclia mapiriensis Encyclia naranjapatensis Encyclia nematocaulon (A.Rich.) Acua Mexico to Nicaragua, and Cuba Encyclia odoratissima Encyclia oncidioides Encyclia ossenbachiana Pupulin Costa Rica Encyclia pachyantha Encyclia patens Encyclia pedra-azulensis Encyclia peraltensis (Ames) Withner Costa Rica Encyclia picta
Size (of the sample) matters Encyclia, are usually less taxonomically problematic. Taxonomy based on morphological analysis still relies on the interpretation of features, but this requires a representation of for species such as Encyclia ceratistes (Fig. 5) or E. mooreana (Fig. 6) allow botanists not only to circumscriptions and correctly interpreting heterotypic synonyms, but also detect those sister species that do 2006; Table 2). A particularly critical situation is posed by taxa national scale, and species comparison has been in many cases limited to the taxa recorded in immediately been critically compared. It is a normal assumption that the patterns of geographic distribution in the masses, and the lack of generalist orchidologists has of Encyclia Bogarn, in preparation), we found one of these cases. specimens for this study, E. amanda was known only TABLE 1. Continues. Species Distribution Encyclia piracanjubensis Encyclia porrecta Encyclia powellii Schltr. Panama Encyclia purpusii Schltr. Mexico Encyclia recurvata Encyclia saltensis Encyclia sclerocladia Encyclia steinbachii Encyclia stellata Encyclia tampensis (Lindl.) Small Florida, Bahamas Encyclia tarumana Encyclia thrombodes Encyclia tonduziana Schltr. Costa Rica Encyclia trachypus Schltr. Ecuador Encyclia triangulifera (Rchb.f.) Acua Cuba Encyclia tripartita Schltr. Mexico to Nicaragua Encyclia wageneri (Rchb.f.) Schltr. Mexico Encyclia yauaperyensis Encyclia yucatanense Schltr. Mexico Epidendrum chloranthm Lindl. Epidendrum halatum Epidendrum glutinosum Epidendrum ramonense Rchb.f. Costa Rica Epidendrum serronianum Epidendrum spectabile LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.330 LANKESTERIANA
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PUPULIN & BOGARN Greenish Encyclia331 FIGURE Encyclia ceratistes. A Bogarn 3800; B Bogarn 3799; C Bogarn 3896 Bogarn 3797; E Bogarn 3798; F Pupulin 5641; G JBL-s.n. Pupulin 5200; I Bogarn 5520; J Bogarn 3806; K Bogarn 3805; L Pupulin 5303; M Bogarn 3803; N Pupulin 5641; O Bogarn 3802. FIGURE Encyclia mooreana. A Bogarn 3810; B JBL-06301; C Bogarn 3721 JBL-08705; E Bogarn 3787; F Karremans 1356; G JBL-08701 Gmez 3; I Bogarn 3792; J JBL-08704; K Bogarn 3790; L Bogarn 3791; M JBL-08708; N JBL-10044; O JBL-08707
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.332 LANKESTERIANAfrom a limited number of specimens. Withner (1998) mentioned no more than eight collections in the distribution with no specimen citation (Withner, 2001). common Costa Rican taxon is not distinguishable from South American specimens of Encyclia chloroleuca (which has nomenclatural priority), and the study of the type specimen of Epidendrum chloroleucum at the basis of the materials at our disposition, it is quite possible that the name E. amanda should be maintained for a distinct species from Panama and, perhaps, northern Colombia. The taxonomy of the Andean and to synonymy with E. chloroleuca. Cleistogamous Encyclia Encyclia, we will accept the concept of E. gravida the name and the real identity of this taxon. Lindley (1849) originally described Epidendrum gravidum on Hartweg s.n., the type!). The holotype at Kew (Fig. 10) consists Oakes Ames for study); the remaining sepals measure 10-11 mm in length and ca. 3 mm in width (Lindley, et al., 1994). Lindley (1849) the original description, the name has been mostly adopted to identify populations of E. stellata, which shares with E. gravida few cases the material referred to seems to match the original concept of Lindley. production of permanently closed, self-pollinated TABLE Encyclia .
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PUPULIN & BOGARN Greenish Encyclia333 FIGURE Encyclia chloroleuca. A Bogarn 2537; B Bogarn 2532; C Bogarn 2537 Bogarn 3111 ; E Bogarn 2544; F Pupulin 3043; G Ossenbach 365; Bogarn 2537; I Bogarn 2544; J Pupulin 3045; K Pupulin 6536; L Pupulin 3045; M Pupulin 3045; N Pupulin 3043; O Pupulin 3044 Spirit. Scale bar = 1 cm. FIGURE Encyclia chloroleuca Pupulin 3045, 11 June 2003 (B), and 3 March 2004 (C). Vouchers at JBL-Spirit. Scale bar = 1 cm. and in the Orchidaceae this is usually accomplished by direct transfer of pollen grains from anther to stigma. be the only type produced, but they may also appear together on the same plant along with typically insect
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.334 LANKESTERIANA Encyclia E. gravida pollinated in the wild, and the pollination rate is high of E. ceratistes are found in pristine and altered forms, the only way to tell the two entities apart is and elongate while the buds are in their early stage of fruit approaches maturity, whereas in allogamous, place but begins to dry out a few days after pollination and becomes papyraceous with age. We found that there is no way to distinguish between the fruit of a turgid, without opening the perianth and looking at the remnant of the rostellum, which is completely absent in cleistogamous forms. This may perhaps explain encyclias is scanty. Before World War II, Blanche Ames (Ames, 1923) later destroyed (Fig. 11, A), which may correspond morphologically to the species originally described by Lindley. A Puerto Rican cleistogamous specimen was documented by Ackerman (1995), who noted that another collection recorded by Schweinfurth (1940) from Maricao in Puerto Rico (, AMES) belongs to this concept, because, according to We had the plants growing under controlled conditions for a while, and cleistogamy in this case appears to be share some of the morphological features expected for this taxon, they also differ in a number of characters, in particular the shape of the petals, the length of the on the midlobe, and the shape of the midlobe, which FIGURE Epiden drum chloroleucumFIGURE 10. The holotype of Epidendrum gravidum. Reproduced with permission by the Board of Trustees, Royal Botanic Gardens, Kew.
the fact that whereas one of the specimens lacks locality data, the other was collected in a region where only one other species of Encyclia, E. ceratistes, occurs naturally. Encyclia specimens in cleistogamous specimen. This may perhaps indicate that essential requirements to be considered a good species, and are nothing more than occasional mutants. When we compare the few documented specimens of cleistogamous Encyclia could attribute these differences to some degree of deformity of the perianth parts, which do not spread E. gravida? largely uniform in Encyclia species, and the features of plants of Encyclia to compare, this key feature alone the records simply represent self-pollinating forms belonging to different taxa cannot be discarded on to E. gravida with those of other sympatric species of Encyclia could represent an important step toward a better circumscription of the taxa in this group and different pollination strategies. A case for barcoding In recent years, sequencing of plant genomes has been regarded as a powerful tool to assess the barcodes can be used as the basis for nomenclatural to describe new species. Species descriptions are One of the necessary steps to place barcoding within the context of a rich taxonomic approach is the analysis species complexes should be sequenced. Application of the same species concept in Encyclia, where with other disciplines such as phylogeography, LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. PUPULIN & BOGARN Greenish Encyclia335 FIGURE 11. Cleistogamous Encyclia A Mexico, Purpus s.n., (B, destroyed); B Puerto Rico; C Costa Rica, Pupulin 6670 Pupulin 5377 (JBL-Spirit). A, drawn by Blanche Ames, from Ames,
ACKNOWLEDGMENTS and Telecommunications (MINAET) and its National collecting permits under which wild species treated in this W are acknowledged for their courtesy and support during LITERA TURE CITED the Virgin Islands. Mem. New York Bot. Gard. 73: iii-203. Schedul. Orch. 4: 1-62. fourth polymorphic alliance in Epidendrum. Bot. Mus. 3: 93-112. The genus Epidendrum in the United States and middle America. Botanical Museum, Cambridge, UK. 9: 59-70. Culley, T. M. and Klooster, M. R. 2007. The cleistogamous 73: 1. Encyclia (Orchidaceae). Brittonia 13: 253-266. complejo Epidendrum. Orqudea (Mexico City) 9: 277298. of Encyclia on the Orchidaceae. IV. Phytologia 21: 433-439. The genus Encyclia Asociacin Mexicana de Orquideologa, Mexico City, Mexico. Prosthechea (Orchidaceae). Phytologia 82: 370-383. 2003. A combined molecular phylogeny of Encyclia (Orchidaceae) and relationships within Laeliinae. Selbyana 24: 165-179. 55: pl. 2831. Lindley, J. 1831. Encyclia Genera and species of orchidaceous plants, 111. Ridgways, London, UK. Lindley, J. 1842. Epidendrum. Edwardss Bot. Reg. 28: 2736. 13. Epidendrum gravidum. 4: 11. Lindley, J. 1853. Folia Orchidacea. Epidendrum. J. Matthews, London, UK. Pupulin, F. 2005. Encyclia Vanishing beauty native Costa Rican orchids 1 (ed. F. Pupuli), pp. 256-267. Rica. Pupulin, F. 2006. Encylia ossenbachiana (Orchidaceae: Laeliinae), a new species from Costa Rica. Selbyana 27: 4-7. Schlechter, R. 1914. Encyclia Die Orchideen pp. 207-212. Paul Parey, Berlin, Germany. (Epidendrum-Stanhopea). Beih. Bot. Centralbl. 36: 421520. 17. 19. Schweinfurth, C. 1940. A notable extension of range of Epidendrum oncidioides gravidum W. M., Soto Arenas, M. A., Culham, A., and Chase, M. W. 2000. A phylogenetic analysis of Laeliinae (Orchidaceae) based on sequence data from internal transcribed spacers 15: 9614. Withner, C. 1996. The cattleyas and their relatives. Vol. 4. The Bahamian and Caribbean species. Timber Press, Portland, Oregon, USA. Withner, C. 1998. The cattleyas and their relatives. Vol. 5. Central America. Timber Press, Portland, Oregon, USA. Withner, C. 2001. Encyclia und sechs wenig bekannte Arten. 52: 444-450.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.336 LANKESTERIANA
as a better indicator of phylogenetic relationships for Kenneth M. Cameron, American Journal of Botany 92: 1025-1032 (2005). The genera Eurystyles Wawra and their consistently epiphytic habit (Fig. 1A-B, 2A-B), in contrast with the ubiquitous terrestrial habit displayed Balogh, 1982; Garay, 1982; Burns-Balogh et al., 1985; et al., 2003). Furthermore, plants in both of these genera consist of small rosettes of lustrous, usually ciliate LANKESTERIANA 11(3): 337. 2011.THE LEAVES GOT IT RIGHT AGAIN: DNA PHYLOGENETICS SUPPORTS A SISTER-GROUP RELATIONSHIP BETWEEN EURYSTYLES AND LANKESTERELLA (ORCHIDACEAE: SPIRANTHINAE) GERARDO A. SALAZAR 1,3 & ROBERT L. DRESSLER 21 2 3 ABSTRACT. morphology, Eurystyles and matK-trnK, trnL-trnF of Eurystyles and two of were included. Both our parsimony and Bayesian phylogenetic analyses Eurystyles and as sister taxa with strong internal support. The RESU M EN. Eurystyles y han sido considerados como slo distantemente relacionados matK-trnK, trnL-trnF) de 29 especies/22 gneros de Spiranthinae (y grupos externos apropiados); tres especies de Eurystyles estructuralmente distintas entre s y dos de fueron incluidas. Tanto nuestro anlisis de parsimonia como el de inferencia bayesiana recobran a Eurystyles y como taxones hermanos con fuerte apoyo interno. El clado Eurystyles/ Eurystyles y KEY WORDS : Orchidaceae, Spiranthinae, molecular phylogeny
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.338 LANKESTERIANA i.e. they are not deciduous as in other Spiranthinae, slender instead of tuberous. Indeed, Eurystyles and always easy to identify to which genus a particular plant belongs (Johnson, 2001; Soto, 1993). structures. On the one hand, in Eurystyles the raceme is condensed (thus appearing capitate) and the numerous whereas in a lax, one-sided raceme bearing a few (usually 1-4) In Eurystyles do not form a distinct, retrorse spur. The base of the Eurystyles subgenus Pseudourystyles which the lip is sessile and lacks basal lobules). The gynostemium is free and elongated, and the rostellum, when present, usually forms a shallowly notched, bear a retrorse spur, sessile labellum gynostemium, and hard, pointed rostellum remnant (Fig. 2C-F). structure between Eurystyles and led instead considered these genera as distantly related, placing them in different generic alliances (Schlechter, 1920; Balogh, 1982; Burns-Balogh et al., 1985) or et al., 2005). A recently published molecular phylogenetic analysis of Spiranthinae based on sequences of nuclear et al., 2006) sustained a sistergroup relationship between Eurystyles and only one species each of Eurystyles and increased numbers of taxa and characters were included in the analysis. In this study, we assess the phylogenetic position of Eurystyles and by conducting cladistic analyses with expanded sampling of both taxa as well as plastid (matK-trnK and trnLtrnF sequence data. Our aims are to clarify the relationships of Eurystyles and with one another and predictors of phylogenetic relationships in these genera Materials and methods Exemplars of three species of Eurystyles, two of and 24 additional species of Spiranthinae (comprising in total 22 genera) of all other subtribes of Cranichideae sensu et al. (2003, 2009) were used as outgroups. A list of the Although we sampled only three of the about 20 species of Eurystyles, they represent much of the SynanthesEurystyles] borealis Burns-Bal., Robinson & Foster (Burns-Balogh et al., the genus encompasses six to eight species (Garay, et al., 2005), all of which seem to be morphological attributes. Molecular methods et al. (2003) and Figueroa et al. (2008). Bidirectional sequence reads
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SALAZAR & DRESSLER Sister-group relationship between Eurystyles and 339 FIGURE 1. Morphology of Eurystyles. A. Plant of Eurystyles cotyledon in situ in a cloud forest of southern Ecuador. B. Flowering plants of E. auriculata (left) and E. cotyledon E. cotyledon from E. cotyledon viewed obliquely from Gynostemium of E. cotyledon from below. H. Apex of previous gynostemium after removal of the pollinarium, showing the broadly notched rostellum remnant. Abbreviations: an = anther; cf = column foot; gy = gynostemium; la = labellum; ne = nectary; ov = ovary; rh = rachis of the raceme; rr = rostellum remnant; st = stigma; vi = viscidium. (B left, from Salazar 7646; B right-H, from Salazar 7642). Photographs by G. A. Salazar.
FIGURE 2. Morphology of Lankesterella A. Lankesterella gnoma in situ in a mata atlntica of southeastern Brazil. B. Flowering plant of Lankesterella ceracifolia side. F. Gynostemium from below. Abbreviations: an = anther; cf = column foot; ds = base of dorsal sepal (or ovary apex?) adnate to gynostemium; gy = gynostemium; la = labellum; sp = spur; st = stigma; vi = viscidium. (B-F from Salazar 7535). Photographs by G. A. Salazar.LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.340 LANKESTERIANA 4.8 (GeneCodes Corp.). Alignment of the sequences sequence similarity (Simmons, 2004). No data were excluded from the analyses due to unambiguous positions were treated as missing data. Cladistic analyses phylogenetic studies of Spiranthinae and other
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SALAZAR & DRESSLER Sister-group relationship between Eurystyles and 341 TABLE 1. Taxa studied, voucher information, and GenBank accessions.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.342 LANKESTERIANA
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SALAZAR & DRESSLER Sister-group relationship between Eurystyles and 343 et al., 2003, 2009; Figueroa et al., support as compared with the separate analyses. sets in combination. The combined matrix was thus maximum parsimony and Bayesian inference. The parsimony analysis was conducted with the computer (Swofford, 2002), and consisted of a heuristic search with 1000 replicates of random taxon addition for the starting trees and tree rearrangements using tree for storage in memory of multiple trees), and all characters were treated as unordered and had equal weights (Fitch, 1971). Internal support for clades was assessed by bootstrapping (Felsenstein, 1985), for which 300 bootstrap replicates were performed, each with 20 replicates with random taxon addition and TBR branch-swapping, keeping up to 20 mostparsimonious trees from each addition replicate. The Bayesian analysis was carried out using the program et al., region, matK pseudogene, trnK intron, trnL intron, and trnL-trnF intergenic spacer) were selected using the Akaike Information Criterion (Akaike, 1974) with the program Modeltest 3.7 (Posada and Crandall, 1998). In all instances, a six-parameter model with among-site rate heterogeneity modeled according to
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.344 LANKESTERIANAcharacters was selected, except that for the trnK character partitions were declared in MrBayes, one including the trnK intron and another encompassing all the other sequence data, specifying the appropriate models. All model parameters were unlinked among et al., 2005). Two simultaneous analyses were run for hundredth generation under the default conditions of generations (2500 trees) of each run were discarded as the burn-in. Inferences about relationships and posterior probabilities of clades (PP) were based on a majority-rule summary tree constructed by pooling the remaining 15,000 trees. Results Parsimony analysis The concatenated data set consisted of 4549 characters, 1066 (23%) of which trees with a length of 4381 steps, consistency index retention index of 0.64. The six cladograms differed only in the resolution among species of Eurystyles and in whether or not the Stenorrhynchos and clades (see below) are sisters with one One of the six trees is shown in Figure 3A. The same tree, which is topologically identical to the Bayesian consensus and on which bootstrap percentages and posterior probabilities (from the Bayesian analysis; see below) were included for the pertinent clades, is depicted in Figure 3B. Four major clades were (BP 100; Fig. 3A, B): 1) the Stenorrhynchos clade (Stenorrhynchos glicensteinii through Sacoila lanceolata; BP 98%); 2) the clade ( Coccineorchis cernua through ; BP 80%); 3) a strongly supported clade consisting of Eurystyles and (BP 100%); and 4) a strongly supported Spiranthes clade (Hapalorchis lineatus through Dichromanthus cinnabarinus; BP 86%). Eurystyles and are in turn Spiranthes clade. Bayesian analysis The majority-rule consensus calculated from 15,000 trees from the Bayesian identical to the parsimony tree of Figure 3A. Posterior probabilities (PP) of clades are displayed in Figure a strongly supported clade consisting of Eurystyles and (PP 1.00). Discussion the same phylogenetic patterns, both supporting Eurystyles and as forming a strongly supported clade. These congruent results are subtending Eurystyles and (in the parsimony tree portrayed Figure 3A, the branch leading to Eurystyles is 124 steps long, whereas that of is 116 steps long). It has been proposed that parsimony may be inconsistent as a method of phylogeny reconstruction when different groups which occurs when long-branched lineages that are of similarities due to independent substitutions to the same base from different ancestral bases (e.g. Lewis, 1998; Sanderson et al., 2000; Anderson & Swofford, 2004; Bergsten, 2005). Methods based on stochastic models of nucleotide substitution such as maximum Swofford et al., 1996; Lewis, 1998; Felsenstein, 2004; Bergsten, 2005). In these analyses, both parsimony and indicating that the strongly supported placement of Eurystyles and as sister groups is not an artefact of long branches misleading parsimony. Our results are in full agreement with the structure between Eurystyles and in
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SALAZAR & DRESSLER Sister-group relationship between Eurystyles and 345 FIGURE 3. Phylogenetic relationships of Eurystyles, Lankesterella and other Spiranthinae inferred from combined nuclear ITS and plastid trnK-matK and trnLtrnF DNA sequences. A. One of the six shortest trees recovered by the parsimony analysis, with branch lengths drawn proportional to the number of character changes supporting them; arrows point to clades that collapse in the strict consensus. B. Same parsimony tree as in A, which was topologically identical to the Bayesian summary tree, and on which bootstrap percentages (numbers above branches) and posterior probabilities from the Bayesian analysis (numbers below branches) are superimposed (see text).upon closer scrutiny the morphological gap between these genera appears not as large as it would appear (1992), labellum structure between Eurystyles subgen. Pseudourystyles and in both genera, and although species of Eurystyles lack Gynostemium morphology between these genera as described in the literature seems to be dissimilar, with the gynostemium of often being et al., 2005) and a prominent column Eurystyles and look similar, with the main difference being that, in the basal portion of the dorsal sepal or Were it not for this adnation, the proportions of column more similar. pollinarium in narrow point that is absent in Eurystyles Eurystyles
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.346 LANKESTERIANAand both Eurystyles and are known for their frequent, seemingly autogamous forms, in which These phenomena indicate that rostellum morphology structures between these two genera are almost certainly related to different pollination mechanisms. Unfortunately, nothing is known about their natural pollination besides the aforementioned recurrence of Eurystyles auriculata and E. cotyledon produce conspicuous, pleasant, diurnal odors, which might play a role in pollinator attraction. Their pendulous, dense clusters of tubular, tube may be pollinated by small insects (possibly in the they are also fragrant or not. epiphytic habit of Eurystyles and on the one hand and their long branches in the molecular tree on the other, one has to ask whether there may be a relationship between epiphytism and an accelerated the branch of Hapalorchis lineata (Fig. 3A), a terrestrial conclusions in this respect is not possible at this time, but it will be interesting to address this matter as more sequences of these and other genera of the subtribe In closing, it is worth mentioning that our phylogenetic study of Malaxideae (from which our epigraph was taken) in showing that at least in some be a better predictor of phylogenetic relationships Malaxideae and the orchidoid subtribe Spiranthinae, that the species are pollinated by different agents, or in different ways, and the species coexist without interbreeding. There may also be parallelisms in Trichocentrum Poepp. & Endl. and other Oncidiinae, with or without oil glands (see Chase et al., 2009 and references included there). Thus, one might expect that ACKNOWLEDGMENTS. We thank the Curators of AMES, USJ, and VEN for courtesies extended during study of their orchid collections; Jim Ackerman, Pilar Candela, material; Jos Portilla and the staff of Ecuagenera and the enjoyable company and expert guidance to G.A.S. during LITERATURE CITED Akaike, H. 1974. A new look at the statistical model 19: 716. Anderson, F. E. & D. L. Swofford. 2004. Should we be worried about long-branch attraction in real data sets? Investigations using metazoan 18S rDNA. Molec. Phylogen. Evol. 33: 440-451. Spiranthinae (Orchidaceae). Am. J. Bot. 69: 1119-1132. Burns-Balogh, P. 1986. Synopsis de la tribu [sic] Spiranthinae en Mxico. Orqudea (Mexico City), n.s. 10: 47-96. Burns-Balogh, P., H. Robinson & M. S. Foster. 1985. The and a new genus from Paraguay. Brittonia 37: 154-162. Bergsten, J. 2005. A review of long-branch attraction. Cladistics 21: 163-193. Cameron, K. M. 2005. Leave it to the leaves: a molecular
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SALAZAR & DRESSLER Sister-group relationship between Eurystyles and 347phylogenetic study of Malaxideae (Epidendroideae, Orchidaceae). Am. J. Bot. 92: 1025-1032. Chase, M. W., N. H. Williams, A. Donisete de Faria, K. M. Neubig, M. do C. E. Amaral & W. M. Whitten. 2009. Floral convergence in Oncidiinae (Cymbidieae; Orchidaceae): an expanded concept of Gomesa and a new genus Nohawilliamsia Ann. Bot. 104: 387-402. Dressler, R. L. 1981. The orchids: natural history and Massachusetts, USA orchid family. Dioscorides Press, Portland, Oregon, USA. Felsenstein, J. 1978. Cases in which parsimony or compatibility methods will be positively misleading. Syst. Zool. 27: 401-410. approach using the bootstrap. Evolution 39: 783-791. Felsenstein, J. 2004. Inferring phylogenies. Sinahuer, Sunderland, Massachusetts, USA. Figueroa, C., G. A. Salazar, A. Zavaleta & M. Engleman, M. 2008. Root character evolution and systematics in Cranichidinae, Prescottiinae and Spiranthinae (Orchidaceae, Cranichideae). Ann. Bot. 101: 509-520. Zool. 20: 406-416. Garay, L. A. 1982. A generic revision of the Spiranthinae. Bot. Mus. Leaf. 28: 277-425. J. Minasiewicz & D. L. Szlachetko. 2006. Phylogenetic relationships within the subtribe Spiranthinae s.l. (Orchidaceae) inferred from the nuclear ITS region. Biodiv. Res. Cons. 1-2: 18-24. Hendy, M. D. & D. Penny. 1989. A framework for the quantitative study of evolutionary trees. Syst. Zool. 38: 297-309. Johnson, A. E. 2001. Las orqudeas del Parque Nacional Iguaz. Literature of Latin America, Buenos Aires, Argentina. Lewis, P. O. 2001. Phylogenetic systematics turns over a new leaf. Trends Ecol. Evol. 16: 30-37. Ronquist, F., J. P. Huelsenbeck & P. van der Mark. 2005. MrBayes 3.1 manual, draft 5/17/2005. Program Documentation and Manual. Website at http:// morphbank.ebc.uu.se/mrbayes/. Salazar, G. A., 2003. Spiranthinae. Pp. 22-23 in: A. M. Pridgeon, P. J. Cribb, M. W. Chase, & F. N. Rasmussen (eds.), Genera Orchidacearum, Volume 3. Orchidoideae part 2, Vanilloideae Oxford University Press, UK. Salazar, G. A. 2005. Eurystyles Wawra. Pp. 320-321 in: F. Pupulin (ed.), Vanishing beauty: native Costa Rican orchids, Vol. 1. Universidad de Costa Rica, San Jos, Costa Rica. Salazar, G. A., M. W. Chase, M. A. Soto & M. Ingrouille. 2003. Phylogenetics of Cranichideae with emphasis on Spiranthinae (Orchidaceae, Orchidoideae): evidence from plastid and nuclear DNA sequences. Am. J. Bot. 90: 777795. Salazar, G. A., L. I. Cabrera, S. Madrin & M. W. Chase. 2009. Phylogenetic relationships of Cranichidinae and Prescottiinae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences. Ann. Bot. 104: 403-416. Soto, M. A. 1993. Eurystyles a new generic record for the (Mexico City), n.s. 13: 269-274. Kahn & S. G. Brady. 2000. Error, bias, and long-branch attraction in data for two chloroplast photosystem genes in seed plants. Molec. Biol. Evol. 17: 782-797. Swofford, D. L. 2002. PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer, Sunderland, Massachusetts, USA. Swofford, D. L., G. J. Olsen, P. J. Waddell & D. M. Hillis. 1996. Phylogenetic inference. Pp. 407-514 in: D. L. Hillis, C. Moritz & B. K. Mable (eds.), Molecular systematics, 2nd edition. Sinhauer, Sunderland, Massachusetts, USA. Szlachetko, D. L. 1992. Notes on Eurystyles (Orchidaceae), with a description of a new species from Mesoamerica. Fragmenta Flor. Geobot. 37: 13-19. Szlachetko, D. L. 1995. Systema orchidalium. Fragmenta Floristica et Geobotanica (Supplement) 3: 1-152. Szlachetko, D. L. & P. Rutkowski. 2000. Gynostemia orchidalium I. Apostasiaceae, Cypripediaceae, Orchidaceae (Thelymitroideae, Orchidoideae, Tropidioideae, Spiranthoideae, Neottioideae, Vanilloideae). Acta Bot. Fennica 169: 1-379. Szlachetko, D. L., P. Rutkowski & J. Mytnik. 2005. Contributions to the taxonomic revision of the subtribes Spiranthinae, Stenorrhynchidinae and Cyclopogoninae (Orchidaceae) in Mesoamerica and the Antilles. Polish Bot. Stud. 20: 3-387.
LANKESTERIANA 11(3): 349. 2011.ORCHID SEED STORES FOR SUSTAINABLE USE: A MODEL FOR FUTURE SEED-BANKING ACTIVITIES PHILIP T SEA TON & HUGH W. PRITCHARD ABSTRACT. 22 countries. In the longer term, our aim is to expand the network to include more institutions from around the situ and in situ Cattleya quadricolor, a Colombian endemic. The story of C. quadricolor is a familiar one: loss of habitat combined with illegal collection of this beautiful species by endangered species, a National Action Plan for Cattleya species, participation in OSSSU, a group of amateur and professional growers with the necessary expertise to germinate seed for the project and produce seedlings, a willing to act as a focal point and to re-introduce plants of C. quadricolor raised by the project into a number of secure sites. RESU M EN. Los Almacenes de Semillas de Orqudeas para Uso Sostenible (Orchid Seed Stores for Sustainable Use OSSSU P O dedicada as como algunos de los suministros consumibles necesarios. Se ha acordado una serie de protocolos, que pases tropicales y templados que cubren una amplia gama de tipos de hbitat en un solo medio de germinacin e in situ queda ilustrado con el caso de Cattleya quadricolor, una endmica colombiana. La historia de C. quadricolor es algo muy familiar: prdida de hbitat combinada con la ilegal recoleccin de esta especie tan hermosa debido a de Cattleya
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.350 LANKESTERIANA strategy. As long ago as 1984 at the Miami World Orchid Conference it was agreed that orchid seed banking had years later, as it is becoming increasingly apparent that in situ techniques are Pritchard, 2001). Adopted in 2002 by the Conference of the Parties the long-term goal of the Global Strategy for Plant through the setting of 16 outcome-oriented targets for in accessible collections, preferably in the country of origin, and 10% of them included in bgci.org/plants2010/t8/). As 2010 approaches, the orchid community progressed in terms of meeting time frame. Orchid Seed Stores for Sustainable Use (OSSSU), Pritchard and Phil Seaton as a project with the modest aim of storing seeds representing 250 orchid species exceeded by a considerable margin. projects are awarded with the aim of assisting OSSSU initially focused on orchid-rich countries in Asia and Latin America. At the beginning of the project, participants from China, India, Indonesia, the Philippines, Singapore, Thailand, and Vietnam were Cuba, Ecuador, and Guatemala attended a workshop at Quito Botanical Gardens, Ecuador (Seaton & Pritchard, hotspots are mainly concentrated in the tropics (Cribb Chile. membership of the orchid community from orchid his or her own experience and expertise to the project. Once a formal agreement had been signed (a Memorandum of Understanding) with a participating to enable each participating country to purchase a -20 C, plus some consumables. Participants were also under less than ideal conditions (Knudson, 1954), at the time of the 1984 World Orchid Conference there semillas, y un jardn botnico que est dispuesto a actuar como punto focal para re-introducir plantas de C. quadricolor KEY WORDS : orchids, seed stores, Cattleya quadricolor
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SEA TON & PRITCHARD Orchid seed stores351remained a lack of numerical data for initial germination percentage enabling comparisons to be made with later samplings of stored seed. Although doubts continue orchid seed in dry storage (Neto & Custodio, 2005), correctly, good-quality seed of many orchid species are limited. One of the many strengths of OSSSU is that 250 species across many genera in many countries and many different habitats around the globe. All seed accessions will be tested on one medium, Knudson C (Knudson, 1946), to allow a direct comparison to be made between the responses of species and genera. In addition, a sample of each seed lot will be sown on a second medium as a comparison. We already know that Knudson C is not the most suitable medium for higher percentage germination on Murashige and Skoog medium (Murashige & Skoog, 1962) than on percentage germination on Knudson C. Some species will perform better on other media (Arditti et al., species. One aim of the project is to publish all of the data at wider orchid community. Each participating institute is committed to promoting the project through a public work as part of OSSSU. It is no exaggeration to say of communication, has made international projects such as OSSSU possible. A key component of any project in the 21st century is the design and setting audience. The OSSSU website can be found at http:// osssu.org. The publication Growing Orchids from Seed was written with the express purpose of making basic as a basic laboratory manual, the methods described require a minimum of laboratory equipment and are therefore suitable for use where resources are limited. in the orchidrich countries of Latin America, OSSSU has sponsored the translation of the information into Cultivo de Orqudeas por Semillas (Seaton & Ramsay, 2009). in this direction, Growing Orchids from Seed has also been translated into Chinese. At the outset of the project two people at each institution were funded to attend the workshops in an training and expertise to a large number of students and permanent staff who are presently engaged in testing are seeking answers to a number of important questions. Seed of some orchid species are undoubtedly shorteret al., 1999), and genera such as the South African Disa (Thornhill & Stanhopea and Coryanthes retain a reputation for should we be considering additional storage at -196 C using liquid nitrogen? collections both for research and educational purposes. In addition, seedlings can potentially be used as part of re-introduction projects. An example of how OSSSU can contribute to
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.352 LANKESTERIANAcurrently underway in Cali, Colombia (for a detailed account of the project see Seaton & Orejuela, 2009). different backgrounds are working together to produce propagate, and re-introduce Cattleya quadricolor Lindl. (syn. C. chocoensis the Asociacin Vallecaucana de Orquideologa, Jardn In response to increasing concern about their status in the wild, an Action Plan has been formulated Cattleya species in Colombia (Niessen & Caldern, 2002). Included in the remaining wild populations, the production of situ collections, and the reintroduction of endangered species. Cattleya quadricolor, a Colombian endemic, www.iucnredlist.org/static/categories_criteria) using the the latest IUCN Red List categories (Calderon, 2007). Although remnant populations remain in dry and transitional humid forests from 600 to current precarious status of the species. In June 2005, a study group from a wide range of backgrounds was created within the Asociacion Vallecaucana de Orquideologa in Cali with the aim Colombian orchid genera. As part of the project they decided to undertake the in vitro production of orchids in danger of disappearing from the wild, including C. quadricolor, with the long-term aim of reintroducing them once more into their natural habitats. The OSSSU project transferred seed storage tubes and some funds to the Jardn Botnico de Cali where dry seed of Colombian orchids, including C. quadricolor, can be stored, both to act as an insurance policy against future losses of wild populations and projects. Germination of stored seed is currently being an OSSSU database. It is anticipated that germination testing will continue beyond the end of the project the aim of reintroducing C. quadricolor to sites that are protected from collectors. from the outset it was not the stated aim of OSSSU collect orchid seed from the wild. The initial focus was cross-pollinating different clones where possible. With the notable exception of some countries such as Colombia, there remains a lack of information about the status of the majority of orchid species in the wild, and few are to be found on the current Global make informed decisions about which orchid species orchid seed collection, it would seem sensible to store data as practicable. The decision to use existing collections was pragmatic. Collecting seed from the wild can be timethe appropriate authorities before embarking on such a of wild-collected material. In Chile, for example, few is indeed focusing on wild-collected seeds. In this symbiotic and asymbiotic germination techniques. the setting up a self-sustaining network of orchid seed banks around the globe for the future and of gathering and collating data on a wide range of orchid knowledge base for future orchid seed bankers and biotechnologists. The project is already expanding to include a number of Associate Members who, at the (Mallorca). Beyond OSSSU, as the network expands,
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. SEA TON & PRITCHARD Orchid seed stores353Conclusion Thus far, OSSSU has facilitated the establishment of a global network of orchid seed banks. In addition to storing seed according to a common protocol, the project has generated data on seed-capsule ripening times, seed numbers per capsule, media preferences for more than 250 species, the performance of more than 250 species on a single growth medium (Knudson orchid seeds representing species from a wide range of habitats. The next step is to publish these data and regenerate plant material from that seed. Our longterm aim is to expand the orchid seed banking network to include additional countries and institutions and establish a Global Orchid Facility that brings together the information garnered from OSSSU with the dedicated website. LITERA TURE CITED Nishimura & R. Ernst, R. 1982. Orchid seed germination and seedling culture a manual. Pp. 243370 in York, USA. Libro rojo de plantas de Colombia. Volumen 6, Orqudeas: primera parte. Colombia. are there? Pp. 161-172 in: A. Raynal-Roques, A. World Orchid Conference. Naturalia Publications, Turriers, France. 26: 155-158. 1996. The state of the worlds plant genetic resources for food and agriculture. FAO, Rome. Knudson, L. 1946. A new nutrient solution for the germination of orchid seed. Amer. Orch. Soc. Bull. 14: 214-217. Amer. Orch. Soc. Bull. 22: 260-261. Pp. 165-181 in Murashige, T. & F. Skoog. 1962. Tissue culture a new means of clonal propagation of orchids. Amer. Orch. Soc. Bull. 33: 473-478. Neto Machado, N. B. & C. Castilho Custodio. 2005. Orchid Selbyana 26: 229-235. Niessen, A. & E. Caldern. (ed.) 2002. Plan de Accin para Cattleya en de Recursos Biolgicos A Boletn No. 30.
LANKESTERIANA 11(3): 355. 2011.FITNESS LANDSCAPES IN ORCHIDS: PARAMETRIC AND NON-PARAMETRICAPPROACHES RA YMOND L. TREMBLA Y 1,2,31 2 3 ABSTRACT. Natural selection and genetic drift are the two processes that can lead to cladogenesis. Without a models, which assume that the parameters are normally distributed. If we forego the idea that selection follows measured characters, whereas non-parametric approaches may be more useful as a tool to detect selection differences among characters. RESU M EN. usualmente puestas a prueba con modelos generales que asumen que los parmetros estn distribuidos en forma Tolumnia variegata, Lepanthes rupestris, y Caladenia valida. KEY WORDS: drift) or non-random processes (natural selection). ability to measure the number of such offspring in the
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.356 LANKESTERIANA orchid would require monitoring orchid seeds, which cannot be seen or followed in the wild (in most cases) genetic markers are used. stigma, number of fruits, number of seeds, length of the life span, etc. It is assumed that the number correlated with the number of seeds produced and Traditionally the models of natural selection et al., 2001). These relationships are used as null models for based on the idea that natural selection follows mathematical equations. Consequently, they require assumptions on how natural selection functions, the most serious of which is that phenotypic and natural selection follow pre-established mathematical equations. In a simple example, let us assume that (fruit set). A model of selection built from a linear equation would predict that selection should result in limitations are likely to be present; perhaps large Box 1. Traditional models of selection for linear ( POSITIVE and negative) selection, stabilizing, and disruptive selection. Linear models test the possibility of either the small or large form of a character having a selection advantage following a linear equation. The quadratic equation for stabilizing selection tests id the medium-size character has an advantage over the small and large sizes. The quadratic equation for disruptive selection tests if small and large sizes have advantages over the intermediate-size character.Phenotypic selection In most cases, phenotypic selection is the process the underlying architecture of genetic inheritance of the character in question is unknown. The assumption is genetically based, which is the case for all orchid studies to date. The complexity of the genetic architecture quickly the character can be inherited among generations selection in the wild with examples from the orchid following discussion into two parts. First I show how the traditional approach (Lande & Arnold, 1983; Endler, 1986), and then I discuss a non-traditional approach to the problem of detecting phenotypic selection in the wild (Schluter, 1988). Common regression approach The traditional approach is described in detail by Lande and Arnold (1983; see also Endler, 1986 for a simpler description of the mathematics). They showed how to use multiple regression analyses of untransformed character traits that are regressed on the
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. TREMBLA Y Fitness landscapes in orchids357 BOX 2. Example of calculations of basic parameters for a hypothetical orchid in a population of 6 individuals assuming T1 = = number of fruits. Note that the mean of the relative will always sum to 1.00.estimates to quantify the strength of selection. The step is estimating the slope of the linear regression (multiplying the trait character as follows: [(T1i-mean T1)2, where T1 and T1i addition, the effect of interaction among traits (T12 = T1 x T2; where T2 will not discuss the effect of interactions among traits in this paper. See Boxes 2 and 3 for examples of how to Selection differentials and selection gradients The between the population mean before and after selection. This should not be confused with a selection gradient, There are three types of indices of used in this case is a logistic regression, where the proportional response is plotted as a consequence of the produced twice as many offspring as the mean. These data are most often plotted as a response to a continuous BOX 3. The regression analysis. In this case I use common statistical software to test for linear disruptive and stabilizing selection The data are from Box l. Analysis performed with the statistical program JMP A selection of the test results shows a partial table with statistical values; a p< 0.05 is considered signi ecology We observe that p = 0.0042 for wer size, so we hypothesis and accept that there is a relationship between wer size and fruit set. The quadratic function i s not sign ere. The shows the relat ionship betwwer size and r. The solid red line shows the mean and stippled lines the 95% l of the line The line is calcu lated (estimated) as -5.208 + 0.213* wer size. Note that the line is the best estimate of the relationship between the two continuous variables.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.358 LANKESTERIANAAssumptions of the regression analysis Regression analysis has a number of assumptions. For example, the of response that follows a normal distribution; 2) across are selected at random; and 5) there is no error in the measurement of x. These are the conditions for testing the quadratic function is the null hypothesis, then the Non-parametric approach The limitation to the parametric approach is that be some other function that does not follow a linear or a cubic spline approach and allow the data to inform approach we do not assume that relationship between (linear, quadratic, etc). This method, which has been Schluter and Nychka (1994), and Tremblay et al. (2010), is a two-step process and can be applied using html). complexity of the equation. A large range of possible of equation are used, OCR and GVC scores (Schluter, (e.g. fruit set). The mathematics behind the application of the cubic spline is complex and not the goal of this paper. Those interested should search the references introduction to the concept. The process of performing the analysis is presented in Boxes 4-7. Caveats of the cubic spline approach The down side of this non-parametric approach using cubic spline for applicable to both methods. The challenge is detecting phenotype. For example, let us consider a hypothetical epiphytic orchid. When the orchid is growing in a section of a tree where the substrate is decomposing, BOX 4. Application of the program GLMS from Dolph Schluter (University of Briti sh Columbia). Step 1: The data are entered in a text saved as a document with the .dat extension. Fitness data (number of fruits) are entered in the character is entered in the second column. Examp le of data entry 0 24 1 25 2 29 3 30 5 33 8 34
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. TREMBLA Y Fitness landscapes in orchids359 BOX 5. Determining the best lambda will determine how complex the line must be to explain the data. Step 2: Starting with a range of lambda from -10 to 10, run the program. Notice that both the GVC and OCV score dip in the range of -2.00, indicating that best lambda to explain the data is in this area. Re-run the program choosing a range of lambd a from -3.00 to -1.00. Notice that minima are now in the range of -2.2 to l.4, depending on which index is evaluated. To evaluate the difference between the two indices, it is recommended that you evaluate both minima. Let us choose -1.6 for the following steps. In general, the differences between the two scores (thus the ranges) have minimal effect on the ultimate outcome. plasticity of expression in orchids can be dramatic as a response to resources, to the point that sex expression Examples of phenotypic selection in orchids Cintrn-Berdecia and Tremblay (2006) showed that selection on column length of Lepanthes rupestris Stimson was likely present in some populations at certain time periods and that there was BOX 6. Constructing the relationship between character variat ion. Step 3: We apply a lambda of -1.6. Then we choose the number of bootstrap replicates (to calculate the error around the best line). Choose a bootstrap of 1000 or more. BOX 7. Step 4: Observation of the ess landscape for the trait under study. Note that the line is not strai ght as predicted by the line ar model. The rate of change as a consequence of size varies across the morphological landscape. Our hypothetic al orchid with a character trait size of 30 has a ess advantage of 1 (the mean of the population), whereas a plant of size 32 has 1.5 as its advantage. In other words, it is expected to have 50% more progeny than a plant of size 30, and an individual with a size of 29 would be expected to have half the ss of an individual of size 30.all populations. They showed that larger columns
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.360 LANKESTERIANA Caladenia valida (Nicholls) M.A.Clem. & states of Victoria and South Australia, usually bears a from the tip of the lateral sepal to tip of the petal, a possible measure of what the pollinator may see from far away. It is also assumed that the tips of the sepals et al., 2006; Faast et al., 2009; Phillips et al., 2009). The same probability of setting fruit, indicating a plateau has been reached with no selection detected among in Tolumnia variegata (Sw.) Braem, a twig epiphyte, is the most widespread species of the genus with a distribution from the Virgin Islands in the eastern and the Cayman Islands. Plants commonly occur on shrubs and small trees mostly in secondary habitats (Ackerman, 1995). Plants at Tortuguero, Puerto Rico, the year (Ackerman, 1995). The orchid is selfincompatible, offers no pollinator rewards, and is Pollinators are female Centris decolorata Lepeletier BOX 8. Comparison of traditiona l and non-traditional approaches advantage in Lepanthes rupestris. Data from Cintrn-Berdecia and Tremblay (2006). The linear relationship between column length and pollinaria removal wm (relative = 197, wm = -.086 + 0.053 Length of the column, r2 = 0.023, p < 0.01. Each point represents an individual, and many individuals overlap. The non-parametric evaluation of the relationship. Short column length has a selective disadvantage, and longer column length is associated with a large increase in the probability of pollinarium removal. The relationship does not follow a simple linear function. Each point represents the mean for the trait of a the interval varies as a consequence of sample size and consistency in the response variable. Squares represent the mean of the response for the
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. TREMBLA Y Fitness landscapes in orchids361 in higher fruit set in this orchid (Sabat & Ackerman, The parametric and non-parametric analysis had similar results (Box 10). The error around the line is smaller for the cubic spline analysis, and the relationship is not linear (although not far from it). In general, the results from these two analyses are similar graph shows the relationship using a logistic regression of size with the probability of fruit on fruit set is detected as p > 0.05. The non-parametric evaluation of the relationship. Short column length has a selective disadvantage, and longer column length is associated with a large increase in the probability of The non-parametric analysis shows that small (<40) is a setting fruit, whereas a of 40-80 has the same expected fruit set. Note the 95% con large for ower size of 50+, and the true trend cannot be predicted with This pattern would be impossible to detect using the traditional approach. Box 10. The linear relation ship in Tolumnia variegata between number of s and relative fruit set. Relative fruit set = 0.599 + 0.0302*# F = 3.557, r2 = 0.01 p= 0.061. Note that the equation explains only 1% of the variation. The cubic spline analysis of number of wers and fruit set. Squares represent the meess for the trait. The relationship is similar to the linear regression but shows a tapering off when plants have many wers.enough to support either method. The cubic spline analysis has the drawback that the estimated means for Discussion of methods difference). The next steps would be to determine
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.362 LANKESTERIANAapproach, the cubic spline results will likely be similar. The limitation of using cubic spline is that no null hypothesis is present, and so a strong a priori bias interpretation of the results. In both approaches, among phenotypes is not likely to be limited to linear and quadratic equations. If we restrict our analyses to only these equations, then we undermine the whole is likely to be more complex than the methods used Caladenia valida can change from one year to the next as a consequence of (Tremblay et al., 2010). pollination syndromes? One of possibilities is that our et al., 2001; et al., 2005). LITERA TURE CITED the Virgin Islands. Mem. New York Bot. Gard. 73: 1-203. Oncidium variegatum: moon phases, pollination and fruit set. Amer. Orch. Soc.Bull. 54: 326-329. intensity and frequency of pollination and the cost of fruiting. Ecology 74: 1033-1042. history of Caladenia. Austral. J. Bot. 57: 247-258. Endler, J. A. 1986. Natural selection in the wild. Princeton syndrome of Caladenia rigida (Orchidaceae). Austral. J. Bot. 57: 315-325. tropical orchids. Oecologia 154: 731-741. & P. Beerli. 2001. The strength of phenotypic selection in natural populations. Amer. Natural. 157: 245-261. Lande, R. and Arnold, S. J. 1983. The measurement of 1226. & R. Peakall. 2009. Implications of pollination by food Caladenia (Orchidaceae). Austral. J. Bot. 57: 287-306. 616-626. from food to sexual deception. Intern. J. Pl. Sc. 167: 1197-1204. Amer. Natural. 143: 597-616. pollinated tropical orchid. Bot. J. Linn. Soc.151: 405410. Riding through the selection landscape: consequences selection landscapes. Philos. Trans. Royal Soc., B 365: 491-498. sex expression in the orchid Ecolog y 72: 5.
PROCEEDINGS OF THE THIRD SCIENTIFIC CONFERENCE ON ANDEAN ORCHIDSPOSTERSCONSERVATION SCIENCE ECOLOGY SYSTEMATICSLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
El presente trabajo pretende desarrollar un sistema protocormos de Oncidium stenotis en nitrgeno lquido (-196 C), con el uso de la tcnica de proyecto consisti en tomar dos fracciones de las semillas, una de ellas fue puesta a germinar en medio y cido giberlico y la segunda fraccin en medio Knudson slido (7 gL-1 de agar) enriquecidos con: actico y cido giberlico. Ambas fracciones se de reguladores de crecimiento en ambas fracciones es encapsular las semillas y protocormos en alginato de sodio al 3%, las cpsulas posteriormente sern deshidratadas en concentraciones crecientes de sacarosa 0.15 M (24 h), 0.25 M (24 h) y 0.5 M (24 h) para luego ser colocadas en slica gel por 5 horas. directa en nitrgeno lquido durante una hora. El por un minuto, luego las cpsulas que contengan semillas se sembrarn directamente y los protocormos slido, con la adicin de diferentes concentraciones buscar el medio adecuado de resuperacin postcongelamiento.LANKESTERIANA 11(3): 365. 2011.APLICACIN DE LA TCNICA DE ENCAPSULACIN DESHIDRATACIN PARA LA CRIOCONSERVACINDE SEMILLAS Y PROTOCORMOS DE ONCIDIUM STENOTIS (ORCHIDACEAE) ALBERTO ROURA 1,3 KARINA PROAO 2 & MNICA JADN 11 2 Carrera en Ingeniera en Biotecnologa, Laboratrio de Biologa Molecular Vegetal, Sangolqu3 Autor para correspondencia: email@example.com CONSERVATION SCIENCELANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
Prosthechea aff. (Mart.) Soto Arenas & epiphytic orchid with a restricted distribution. It occurs in deciduous oak forests, particularly on Quercus deserticola in the northern portion of collected from their natural populations. Learning important, as we may gain knowledge about the genetics and ecology. This will be a useful tool for of P. aff. 2) determine if the species requires an external pollinating agent for sexual success of this species. Exclusions (150) were we applied four pollination treatments: manual Preliminary results showed that outcrossing and set than spontaneous autogamy and open pollination treatments, meaning that P. aff. has Prosthechea aff. is pollinator-dependent for sexual reproduction, taking into account that the spontaneous autogamy treatment showed nil fruit of hand pollination treatments. This indicates that this species, as other species of tropical orchids, is pollination-limited.LANKESTERIANA 11(3): 366. 2011.MATING SYSTEM AND FEMALE REPRODUCTIVE SUCCESS OF THE ENDEMIC, EPIPHYTIC PROSTHECHEA AFF KARWINSKII (ORCHIDACEAE) ERIKA CAMACHO-DOMNGUEZ & IRENE VILADAZ Corrersponding author: firstname.lastname@example.org ECOLOGYLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011. con la reproduccin in vitro de Laelia speciosa el papel de su presencia en la planta secuenciamos con los primers ITS1F e ITS4. Posteriormente se clonaron los fragmentos con TOPO TA4 y terminator en un ABI 3100. En total se secuenciaron correspondieron a secuencias de L. speciosa, aunque hongos. El resto de las secuencias agrupadas en 97% de similitud correspondieron a 19 especies de hongos de los gneros Alternaria, Cylindrocarpon, Curvularia, Fusarium, Myrmecridium, Neonectria, Penicillium, y Tetracladium, as como a especies Nectriaceae, Pucciniomycotina, Sordariomycetes, y Tricholomataceae. Muchos de estos taxa estn presentes en las semillas, lo que sugiere que estos hongos parsitos, causan la muerte de las semillas en L. speciosa es considerable y la biologa de los HONGOS ENDFITOS DE LA ORQUDEA EPFITA LAELIA SPECIOSA ROBERTO GARIBA Y-ORIJEL 1 KEN OY AMA 2 & IRENE VILA-DAZ 3,41 Instituto de Biologa, UNAM23 Michoacn, Mxico4LANKESTERIANA 11(3): 367. 2011.
(1862) as a strategy to promote cross-pollination. labellum in Spiranthes of the column in Sauroglossum elatum Lindl. and Manniella column in Prescottia stachyodes (Sw.) Lindl. In the Aa erosa (Rchb.f.) Schltr. and the mechanism responsible for Aa erosa is straight. The gradual cell death of the dorsal side of side cause the column to bend downward to almost suggested self-pollination in Aa Aa Aa erosa could discard selfpollination as the only strategy of pollination in this genus.EVIDENCE OF PROTANDRY IN AA RCHB.F (ORCHIDACEAE, CRANICHIDEAE) DELSY TRUJILLO THASSILO FRANKE & REINHARD AGERER Corresponding author: email@example.com LANKESTERIANA 11(3): 368. 2011. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
Pleurothallis been proposed. Luer has proposed that Pleurothallis species in subgenus Scopula be segregated into the genera Colombiana and Ancipitia and Margonska (2001) proposed the genus Zosterophyllanthos for Pleurothallis subsection Macrophyllae-Fasciculatae Luer (2005) proposed the genus Acronia by uniting Pleurothallis subsection Macrophyllae-Fasciculatae with subsections Acroniae and Amphygiae. The molecular phylogenetic studies by Pridgeon and here a more detailed phylogenetic analysis of the genus Pleurothallis, with emphasis on subsection Macrophyllae-Fasciculatae, with data primarily from nuclear ITS sequencing, supplemented with conclusions can be drawn. In the strict consensus maximum-parsimony tree of ITS data, many of the used to delimit the genus Pleurothallis. Such a tree would argue for an expanded concept of the genus Pleurothallis, in which the groups Ancipitia, Colombiana, and Acronia/Zosterophyllanthos, if shown to be monophyletic, are relegated to subgenera.LANKESTERIANA 11(3): 369. 2011.A PHYLOGENETIC ANALYSIS OF THE GENUS PLEUROTHALLIS, WITH EMPHASIS ON PLEUROTHALLIS SUBSECTION MACROPHYLLAEF ASCICULATAE, USING NUCLEAR ITS AND CHLOROPLAST DNA SEQUENCING M. WILSON C. BELLE, A. DANG, P. HANNAN, C. KENYON, H. LOW, T ST A YTON & M. WOOLLEY Corresponding author: firstname.lastname@example.org SYSTEMATICSLANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
species within Brachycladium nummularium (syn. Oreophilus nummularius). Sequencing of additional Brachycladium B. stalactites Amplectentes. Since the relationship of Brachycladium to Andinia had been noted earlier, a number of Andinia species clade containing A. lappacea, A. pensilis, and A. vestigipetala that was closely related to Brachycladium and a second, more distantly related clade containing A. dalstroemii, A. pogonion, and A. schizopogon. Samples of two additional genera, Masdevalliantha Xenosia Luer, were included in the analysis based upon morphological similarity to Andinia. These two genera formed a clade that was sister to both Brachycladium and Andinia. Only the node subtending clades of all whereas nodes subtending clades of Brachycladium plus Andinia, or Brachycladium alone, had much lower support. If future chloroplast sequencing produces phylogenies congruent with nrITS, this would indicate that Andinia, Brachycladium (syn. Oreophilus), Masdevalliantha, and Xenosia should all be placed in the same genus.PHYLOGENETIC ANALYSIS OF THE ANDEAN GENUS BRACHYCLADIUM LUER (SYN. OREOPHILUS HIGGINS & ARCHILA) AND CLOSELY RELATED GENERA BASED ON NUCLEAR ITS SEQUENCING MARK WILSON 1,3 & LOU JOST 212 Via a Runtun, Baos, Tungurahua, Ecuador3 Corresponding author: email@example.com LANKESTERIANA 11(3): 370. 2011. LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.
Pleurothallis subsection Macrophyllae-Fasciculatae (syn. Acronia this genus in Mesoamerica and the Andes. To this end, College with plants from commercial operations in South America (Ecuagenera; Colomborqudeas; taken; in the future, herbarium sheets will be prepared or from some locations such as Central America which is approximately 46% of the described species. A genetic barcode for this subsection of Pleurothallis will likely consist of three sequences. While only nrITS has been sequenced for these plants so far, the chloroplast sequences rpoB2, rpoC1 ycf1 to combine with matK or trnH-psbA as per the CBOL Plant Working Group.LANKESTERIANA 11(3): 371. 2011.BARCODING THE SPECIES OF PLEUROTHALLIS SUBSECTION MACROPHYLLAE-F ASCICULATAE MARK WILSON firstname.lastname@example.org LANKESTERIANA 11(3), December 2011. Universidad de Costa Rica, 2011.