THE ORCHID POLLINARIA COLLECTION AT LANKESTER BOTANICAL GARDEN, UNIVERSITY OF COSTA RICA

Adam Karremans

We continue to document the morphology of the pollinaria of the orchids of southeast Mexico, this time concentrating on 56 species from the Tacaná-Boquerón biological corridor, a recognized biodiversity ''hotspot.'' This study is a component part of the project ''Ecology and sustainable cultivation of orchids of Soconusco,'' which aims to conserve and restore the orchid populations of this biodiverse region.

This is the second volume of the series on Orchidaceae for Flora Costaricensis, after the fundamental contribution by Atwood and Mora de Retana, published 10 years ago. The present work was conceived and realized at Lankester Botanical Garden, University of Costa Rica (LBG), as part of the commitment by the center to carry out a complete inventory of the orchid flora of the country and to provide relevant information for the conservation of endangered taxa. In the past 10 years, the activities of botanical exploration carried out by the staff of LBG significantly improved the living plant collections grown at the Garden, revealed several floristic novelties, and made important steps toward a better understanding of species identity in this large and taxonomically complex group of plants. [...] Although they only represent a small portion (around four percent) of the Costa Rican orchid flora, species of Zygopetalinae are frequently grown (and often avidly collected) for horticultural purposes, and their study is crucial to understanding distribution patterns and species frequency as the first step to establish conservation priorities. Among the genera of Zygopetalinae, Dichaea constitutes a particularly common element in any type of vegetation in the country, and its taxonomic treatment should have a certain utility to field botanists working in Costa Rica. Sixteen genera (including a natural hybrid genus) and 60 species are treated.

We investigated the phylogenetic placement of Cyanaeorchis and selected representatives of the tribe Cymbidieae based on nuclear (ITS) and plastid (matK–trnK and rbcL) DNA sequences. Bayesian and parsimony analyses of separate and combined datasets were largely congruent with each other and showed that the Neotropical Cyanaeorchis does not belongs in the predominantly Old World subtribe Eulophiinae, where it has previously been placed. Instead, it is strongly supported as a sister to Grobya in Catasetinae. Because Catasetinae are Neotropical and there are no unequivocal morphological similarities between Cyanaeorchis and other genera in the subtribe, this relationship reflects a geographical rather than morphological similarity and suggest habitat-driven local diversification. Specimens from central Brazil formerly identified as Cyanaeorchis minor are shown to be a distinct species, described here as C. praetermissa. Niche modeling indicates that C. praetermissa and C. minor have di...

Selbyana 29(1): 69- 86. 2008. THE ORCHID POLLINARIA COLLECTION AT LANKESTER BOTANICAL GARDEN, UNIVERSITY OF COSTA RICA FRANCO PUPULIN* Lankester Botanical Garden, University of Costa Rica. P.O. Box 1031-7050 Cartago, Costa Rica, . CA Angel Andreetta Research Center on Andean Orchids, University Alfredo Pérez Guerrero, Extension Gualaceo, Ecuador Harvard University Herbaria, Cambridge, MA, USA The Marie Selby Botanical Gardens, Sarasota, FL, USA Email: fpupulin@cariari.ucr.ac.cr ADAM KARREMANS Lankester Botanical Garden, University of Costa Rica. P.O. Box 1031-7050 Cartago, Costa Rica, CA Angel Andreetta Research Center on Andean Orchids, University Alfredo Pérez Guerrero, Extension Gualaceo, Ecuador ABSTRACT. The relevance of pollinaria study in orchid systematics and reproductive biology is summarized. The Orchid Pollinaria Collection and the associate database of Lankester Botanical Garden, University of Costa Rica, are presented. The collection includes 496 pollinaria, bèlonging to 312 species in 94 genera, with particular emphasis on Neotropical taxa of the tribe Cymbidieae (Epidendroideae). The associated database includes digital images of the pollinaria and is progressively made available to the general public through EPIDENDRA, the online taxonomic and nomenclatural database of Lankester Botanical Garden. Examples are given of the use of the pollinaria collection by researchers of the Center in a broad range of systematic applications. Key words: Orchid pollinaria, systematic botany, pollination biology, orchid pollinaria collection, Lankester Botanical Garden, Universidad de Costa Rica In a family whose ftowers are pollinated mostly by insects, powdery pollen (which is The cohesion of pollen into compact units more suited for the placement on hairy parts) (the pollinia) and the evolution, in the Orchida- was less effective in promoting dispersal, and ceae, of accessory structures for attachment to the evolution of an adhesive viscidium was crupollinators, has attracted the attention of scien- cial to maxirnize the efficiency of pollen export tists since the time of Darwin, whofound orchid through the improvement of its adherence to poIlinaria to be "as perfect as any of the most smooth surfaces such as an insect's thorax, beautiful adaptations in the animal kingdom" mouthparts, frons, eyes, and legs (Dressler (Darwin 1877). If the grouping of pollen grains 1981). The efficiency of pollen transfer depends into tetrads, reduction in exine surrounding inner not only on the selection of visitors by means of grains, and pollen consolidation into pollinia are specific visual and olfactory cues, but also on also known in the Asclepiadaceae, only the Or- the precise deposition of the pollen masses on a chidaceae show the innovation of pollinaria . specific target area of the pollinator's body. This This term refers to the entire package, including • involves modifications in perianth parts as weIl one or more pollinia and the associate structures, as in gynoecial and androecial organs, and evenwhich is removed by pollinators, and its evolu- tually the shape and size of the viscidium. The tion is a key to explain sorne of the major mor- precision of pollen placement on specific parts phological differences between orchids' ftowers of the body of pollinators ensures interspecific pollination and, on the other side, allows several and those of other plant farnilies. sympatric orchids to share the same pollinators without interference in pollen deposition (Dress* Corresponding author. 1er 1968, Nilsson 1983, Steiner 1989). MechanINTRODUCTION 69 70 SELBYANA Volume 29(1) 2008 Pollinari a in dorsal and ventral view s. A. Acineta densa (JBL-09182). B. unide ntified Acineta species (Pupulin 6291). Scale bar = 5 FIGURE 2. PoUinaria in dorsal, ventral, and late ral views. A. Kefersteinia parvilabris (Bogarfn 806). B. Chondroscaphe atrilinguis (Bogarfn 2950). Scale bars = 1 mm. ical movements of the caudicles and stipe after pollinarium removal re-orient the pollinia and maximize pollen distribution. Bending movements of pollinaria, which correctly orient the pollinia to contact the stigma (Darwin 1877, Nilsson 1980, Johnson & Nilsson 1999), as weIl as the shrinking of the pollinia to fit the stigmatic entrance (Borba & Semir 1999), prevent geitonogamy and promote effective outcrossing in pollen transfer. The high degree of variation in pollinaria organization and morphology reflects the central role of this structure in orchid pollination mechanisms, and Johnson and Edwards (2000) suggested it was pivotaI in promoting the tremendous radiation of the Orchidaceae, which have more than 25 ,000 species (Atwood 1986, Cribb & Govaerts 2005). The characteristics of pollinia and the associated structures have been used as a tool for orchid classification since the early work of John Lindley ; Reichenbach (1852) devoted his doc- toral dissertation to the study of poIlinaria as indicators of species relationships. Diversity in number, shape, orientation, and packaging of pollinia (including the degree of tetrad packing, the state of the pollini a, the form, arrangement, orientation, and possible dimorphism of massulae), as well as the structure, development, organization and shape of poIlinaria (including the presence of caudicles and their composition , a nd the presence, number and shape of stipes), have been employed as sources of characters at various systematic levels to define taxa from species to subfamilies. The use of pollinaria information as a n addition al tool in taxonomic work is a current practice at Lankester Botanical Garden (LBG), where researchers make use of their morphological features in a broad range of systematic applications (i.e., Dressler & Klikunas 2006, Pupulin 2007a, 2007b; Bogarfn 2007, Pupulin in press). As pollinaria are less subject to parallelism than other floral features, their morphology also played a significant role in the FiGURE 1. mm. PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 71 and the lack of direct records inhibits a better understanding of the pollination biology of m a n y orchid taxa, and particularly of the real degree of pollinator specificity. The study of mating patterns and male fitness in orchid reproduction is dependent on a precise identification of pollinaria. Pollinaria found on insects are often informative of the presence of rare species or of orchids that occupy difficult to reach habitats and ecological niches. During a preliminary census of the orchid flora of the large «400 hectares) Alberto M . Brenes Biological Reserve in Costa Rica, staff of LBG managed to record the presence of Houlletia tigrina on the basis of a single pollinarium found on an insect that had been captured at the Reserve during a course on entomology. Dressler (1976) makes an interesting case of the study of orchid pollination without any orchids, through the identification of orchid genera and species from the pollinaria borne by captured insects, stressing the importance of a reference collection to compare pollinaria. Recent work by Widmer and collaborators addressed the problem of the relative paucity of field observations by means of a molecular approach, based on the analysis of DNA recovered from pollinaria found on insects (Widmer et al. 2000). The study demonstrates that the analysis of the nuclear ribosomal ITS region of the recovered pollen allows the identification of the species or species-group from which the pollinaria originated, disclosing previously unrecorded orchid-pollinator relationships. ORCHID POLLINARIA COLLECTION AT LANKESTER BOTANICAL GARDEN FIGURE 3. A. Cases hosting the collection of pollinaria. B. E nvelopes to store pollinaria in the collection of Lankester Botanical Garden. construction of phylogenetic hypotheses (i.e., Williams 1970a, 1970b, 1972; Ackerman & Williams 1981 , Burns- Balogh 1982, Dressler 1986, 1993; Chase 1987, Freudenstein & Rasmussen 1997). Accumulating evidence shows that floral evolution in the Orchidaceae is essentially driven by pollinator traits , linking the study of pollination to evolutionary biology. New, intricate orchidpollinator interactions are continuously revealed through the application of accurate empirical studies that improve our understanding of plant evolution in general. However, direct observation of pollination acts is notoriously difficult, To support studies in system atic botany and pollination biology, LBG began in 2003 an effort to build and maintain a large collection of orchid pollinaria, with emphasis on Neotropical taxa and special groups of interest to the research staff. Pollinaria are mostly removed from plants grown in the living collection of the center, inc1uding more than 15 ,000 accessions belonging to approximately 1300 species. Priority was given to Neotropical and Mesoamerican genera and species, and to the groups that exhibit potentially informative pollinarium morphology, with emphasis on taxa with highly specialized reproductive biology-" It is not insignificant that the mostly Neotropical advanced Cymbidieae account for almost 88% of the entire collection. Unlike the generalized use of mounting pollinaria on small paper triangles , to which they are attached by the natural glue of the viscidium, the pollinaria intended for the collection are removed with a thin-pointed instrument, trying to mini- TABLE 1. No. P0340 P0359 P0347 P0362 P0303 P0358 POl17 P0118 P0119 P0120 P0467 P0403 P0086 P0278 P0171 POO13 P0121 P0336 P0071 P0087 P0122 P0246 P0405 P0194 P0224 P0411 P0153 P0045 P0123 P0450 P0456 P0149 POl86 P0404 P0468 P0021 P0088 P0188 P0261 P0150 P0170 POI64 POO05 P0386 P0302 POOOI P0354 P0124 P0381 P0287 P0308 P0044 P0042 P0282 P0481 P0380 P0376 P0409 P0465 P0396 P0339 Volume 29(1) 2008 SELBYANA 72 Orchid pollinaria in the collection of L ankester Botanical Garden, University of Costa Rica. Genus/species Acineta Acineta Acineta A cineta densa Acineta densa Acineta densa Acineta densa Acineta densa Acineta densa Acineta densa Ada aurantiaca Ada aurantiaca Ada chlorops Aerides falcata Aerides multifiora Angraecum eburneum Ang raecum eichlerianum Arpophyllum giganteum Ascocentrum curvifolium Aspasia epidendroides Barkeria lindleyana Batemania leferenzii Bensteinia ramonensis Bensteinia ramonensis Benzingia cornuta Benzingia reichenbachiana Benzingia caudata Benzingia reichenbachiana Brassavola nodosa Brassia arcuigera Brassia arcuigera Brassia arcuigera Brassia a rcuigera Brassia arcuigera Brassia caudata Brassia caudata Brassia gireoudiana Brassia gireoudiana Brassia suavissima Brassia verrucosa Brassia caudata Brassia caudata Brassia arcuigera Bulbophyllum lobbii Catasetum maculatum Catasetum maculatum Catasetum maculatum Cattleya intermedia Chaubardia heteroclita Chaubardiella pubescens Chaubardiella subquadrata Chaubardiella pacuarensis Chaubardiella pacuarensis Chondrorhyncha thienii Chondroryncha thienii Chondroscaphe amabilis Chondroscaphe atrilinguis Chondroscaphe atrilinguis Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor . Country Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Colombia Colombia Costa Rica Thailand Thailand Madagascar Africa Costa Rica Thailand Costa Rica Costa Rica Peru Costa Rica Ecuador Ecuador Costa Rica Ecuador Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Jamaica Costa Rica Thailand Costa Rica Costa Rica Costa Rica Brazil Peru Costa Rica Costa Rica Costa Rica Costa Rica Colombia Ecuador Ecuador Panama Panama Costa Rica Costa Rica Costa Rica Collector Pupulin 6291 G6mez 7 Pupulin 4200 Pupulin 4200 Vargas s.n. Pupulin 6224 Bogarin 1923 Bogarin 1923 Pupulin 5989 Bogarin J 890 Pupulin 5989 Whitten 2052 Bogarin Bogarin Pupulin Pupulin Bogarin Bogarin 2416 1640 4886 5646 2416 2554 Bogarin 2855 Herra J Hamlyn s.n. Pupulin s. n. Bogarin 2609 Elizondo s.n. Rojas 137 Pupulin 6518 Pupulin 6240 Blanco 1072 Pupulin 6281 Pupulin 6513 Bogarin 2950 Bogarin 2950 Pupulin 3508 B ogarin 1894 D ate 14.11.2006 14.11.2006 14.11.2006 14.11.2006 13.09.2006 14.11.2006 09.06.2005 09.06.2005 09.06.2005 09.06.2005 20.07.2007 19.07.2007 13.05.2005 01.09.2006 07 .07.2005 15.12.2003 09.06.2005 26.10.2006 13.05.2005 13.05.2005 09.06.2005 13.06.2006 19.07.2007 17.03.2006 16.05.2006 19.07.2007 19.04.2006 13.05.2005 09.06.2005 20.07.2007 20.07.2007 14.06.2005 22.12.2005 19.07.2007 20.07.2007 22.10.2005 13.05.2005 22.1 2.2005 30.06.2006 14.06.2005 07.07.2005 07.07 .2005 06.08.2003 07.03.2006 13.09.2006 20.07 .2004 14.11.2006 09.06.2005 05.03.2006 05 .09 .2006 20.09.2006 13.05.2005 13.05.2005 05.09.2006 19.08.2007 05.03.2006 12.03.2006 19.07.2007 20.07.2007 12.03.2006 14.11.2006 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER TABLE No. P0341 P0326 P0408 P0410 P0412 P0413 P0429 P0355 P0050 P0049 P0048 P0416 P0168 P0346 P0394 P0342 P0473 P0439 P0379 P0464 P0329 P0469 P0378 P0425 P0325 P0398 P0322 POllO P0047 P0125 P0156 P0067 P0126 P0185 P0475 P0089 P0310 P0352 P0026 P0216 P0301 P0479 P0300 POO03 P0478 P0428 P0388 P0286 P0037 P0029 P0030 P0069 P0272 P0275 P0038 P0274 P0031 P0032 P0281 P0033 P0163 l. 73 Continued. Genus/species Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe bicolor Chondroscaphe eburnea Chondroscaphe eburnea Chondroscaphe embreei Chondroscaphe embreei Chondroscaphe escobariana Chondroscaphe flaveola Chondroscaphe flaveola Chondroscaphe flaveola Chondroscaphe yamilethiae Chondroscaphe yamilethiae Chondroscaphe yamilethiae Chondroscaphe yamilethiae Chondroscaphe yamilethiae Coccineorchis standleyi Cochleanthes aromatica Cochleanthes aromatica Coelia bella Coeliopsis hyacinthosma Coeliopsis hyacinthosma Comparettia falcata Cycnoches amparoanum Cymbidium iridioides Cymbidium iridoiides Cyrtopodium macrobulbon Daiotyla albicans Daiotyla albicans Daiotyla crassa Daiotyla crassa Daiotyla c rassa Daiotyla xanthina D endrobium antennatum Dichaea Dichaea acroblephara Dichaea acroblephara Dichaea acroblephara Dichaea amparoana Dichaea c ryptarrhena Dichaea elliptica Dichaea elliptica Dichaea fra g rantissima subsp. eburnea Dichaea fragrantissima subsp. eburnea Dichaea fragrantissima subsp. eburnea Dichaea globosa Dichaea globosa Dichaea globosa Country Costa Rica Panama Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Panama Panama Ecuador Ecuador Ecuador Ecuador Ecuador Ecuador Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Guatemala (?) Costa Rica Costa Rica Costa Rica Costa Rica India India Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Colombia New Guinea Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Collector Bogar[n 2203 Bogar[n 1914 Bogar[n 996 Bogar[n 426 Bogarin 2874 Bogarin 984 Hoffman s.n. Pupulin 3525 Pupulin 3525 Bianchi 68 Bianchi 68 Bogarin 996 Karremans 883 Dressle r 6826 Dressle r 6826 Pupulin 6510 Pupulin 6510 Pupulin 6289 Pupulin 6474 Pupulin 6474 Pupulin 6474 Pupulin 4637 Pupulin 4637 Pupulin 4701 Pupulin 4638 Pupulin 4637 Pupulin 4229 Pupulin 4477 Pupulin 4377 A guilar J Rojas 114 Pupulin 6416 Salazar Pupulin Pupulin Pupulin Pupulin Pupulin Pupulin s.n. 4372 4372 4257 4257 4257 5891 Pupulin 5049 Bogar{n 1009 Pupulin 4945 Pupulin 5049 Bogar[n 6 79 Pupulin 5662 ｾｵｰｬｩｮ＠ 5133 Pupulin 5133 Blanco 513 Blanco 513 Pupulin 4601 Pupulin 4517 Pupulin 4517 Klark 286 Date 14.11.2006 03.10.2006 19.07.2007 19.07.2007 19.07.2007 19.07.2007 19.07.2007 14.11.2006 13.05.2005 13.05.2005 13.05.2005 19.07.2007 07.07.2005 14.11.2006 12.03.2006 14.11.2006 19.08.2007 20.07.2007 05.03.2006 20.07.2007 17.10.2006 20.07.2007 05.03.2006 19.07.2007 03.10.2006 12.03.2006 03.10.2006 24.05.2003 13.05.2005 09.06.2005 16.05.2005 13.05.2005 09.06.2005 22.12.2005 19.08.2007 13.05.2005 20.09.2006 14.11.2006 22.10.2005 03.05.2006 07.09.2006 19.08.2007 07.09.2006 18.03.2004 19.08.2007 19.07.2007 07.03.2006 05.09.2006 13.05.2005 13.05.2005 13.05.2005 13.05.2005 29.08.2006 0l.09.2006 13.05.2005 0l.09.2006 13.05.2005 13.05.2005 05.09.2006 13.05.2005 07.07.2005 74 TABLE 1. No. P0027 P0284 POO1 2 P0036 P0035 P0040 P0039 P0406 P0271 POl 74 P0041 P0I59 P022J P0390 P0222 pOln P0025 P0043 P0046 P01 32 POO08 P0I 3 1 POl 30 POJ27 P0488 P0489 POO06 P02 18 P0480 P0307 P0299 POl 66 POl54 P041 9 POliS P0060 POI 96 POl 95 P0447 P0219 P0228 P046 1 P03 J9 P0255 P0317 P03 12 P0436 P0442 P0227 P03 13 POOlS P0226 P03 16 P03 11 P0229 P0332 POO17 P0493 POO14 POJ05 P0324 SELBYANA Volume 29(1) 2008 Continued . Genus/species Dichaea morrisii Dichaea obovatipetala Dichaea obo vatipetala Dichaea panamensis Dichaea panamensis Dichaea poicillantha Dichaea poicillantha Dichaea poicillantha Dichaea trulla Dichaea similis Dichaea similis Dichaea trulla Dressleria d ilecta Encyclia ambig ua Encyclia cordigera Encyclia ceratistes Epidendrum Epidendrum Epidendrum Epidendrum centradenia EpidendrL/m odontochilwn Epidendrum schumannianL/m Epidendrum wallisii Epigeneium cymbidioides Eriopsis wercklei Eriopsis werckle i E rycina crista-galli Erythrodes tridax Euryblema anatonwn Euryblema andreae Euryblema andreae Eurychone rothschildiana Galeandra lac us tris Galeottia g randiflora Galeottia grandiflo ra Galeottia grandiflora Gastrochilus calceola ris GastrochilL/S f ormosanum Gongora Gongora Gongora Gongora Gongo ra Gongora Gongora Gongo ra arm eniaca Gongo ra arm eniaca Gongora claviodora Gongora galeata Gongora galeata Gongora leucochila Gongora truncata Gongora unicolor Gongo ra unicolor Govenia ciliilabia Grammatophyllum multiflo rum GrammatophyliL/m scriptum Guarianthe skinneri Haraella retrocalla H eterotaxis sessilis Houlletia tigrina Country Costa Ri ca Costa Rica Costa Ri ca Costa Ri ca Costa Ri ca Costa Rica Costa Rica Nicaragua Costa Ri ca Costa Ri ca Costa Ri ca Costa Rica Costa R ica M exico Costa Rica Costa Rica Costa Ri ca Costa Ri ca Costa Rica Costa Ri ca Costa Rica Costa Rica Costa Rica Phi lippines Costa Ri ca Costa Ri ca Costa Ri ca Costa Rica Panama Colombi a Colombi a Africa Ecuador Costa Rica Costa Rica Costa Ri ca Thailand Asia Costa Rica Costa Rica Costa Ri ca Costa Rica Costa Ri ca Costa Ri ca Costa Ri ca Costa Rica Costa Rica Costa R ica Ecuador M exico Costa Rica Ve nezue la Costa Rica Costa Rica Costa Rica Philippines Phi lippines Costa Rica Taiwan Costa Rica Costa Rica Co llector Whitten 2 1 71 Pupulin 5059 Pupulin 5059 Bogar[n 1075 Karremans 62 1 Bogar[n 187 Pupulin 4662 Ka rremans 1140 Pupulin 5130 Pupulin 5644 Pupulin 4089 Blanco 2812 Pupulin 5156 Pupulin 5520 Pupulin 5570 M ontealeg re 114 PupL/lin 4864 PL/pL/lin 3832 PL/pL/lin s.n. Bogar[n 993 PL/pU lin 6282 PL/pulin 6282 Whitten 2138 Whitten 2058 Blanco 2609 Blanco 2598 Gômez 60 R oj as 111 Pupulin 5214 Pupulin 5758 Serrano 125 Pupulin 4954 PL/pulin 621 7 Clark 209 Bogar[n 2480 Date 04.12.2003 05.09 .2006 13.01.2004 13.05.2005 13.05.2005 13.05.2005 13.05.2005 19.07.2007 29.08.2006 17.07.2005 13.05.2005 07.07.2005 09.05.2006 07 .03.2006 12.05.2006 17.07 .2005 22.12.2005 22.12.2005 22.12.2005 09.06.2005 22.12.2005 09.06.2005 09 .06.2005 09.06.2005 14.01.2008 14.01.2008 25 .02.2005 03.05 .2006 19.08.2007 13.09.2006 06.09.2006 07 .07.2005 14.06.2005 19.07 .2007 0 8.05.2004 13.05.2005 17.03.2006 17.03.2006 20.07.2007 03.05.2006 17.05.2006 20.07 .2007 29.09.2006 2 1.06.2006 29.09.2006 29.09.2006 20.07.2007 20.07.2007 17.05.2006 29.09.2006 19.01.2004 17.05.2006 29.09.2006 29.09.2006 06.06.2006 26.10.2006 12.05.2004 05.02.2008 30.10.2002 J9.05. 2005 03 .10.2006 75 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER T ABLE No. 1. Continued. Genus/species P0051 P0402 POO92 P0200 P0197 P0395 P0417 P0056 P0052 P0426 P0385 P0155 P0057 P0055 POl98 P023 1 P0427 P0193 POO04 P0420 P0054 P0053 P044 1 P0165 P0128 P0090 P0091 P0184 POl 87 P0217 P0393 P0454 P0462 P0444 HuntLeya burtii Huntleya burtii Hygroehilus parishii Kefersteinia Kefe rsteinia andreettae Kefersteinia aurieuLata Kefersteinia eostarieensis Kefersteinia eostarieensis Kefersteinia exeentriea Kefersteinia Laminata Kefersteinia orbieuLaris Keferste inia orbieuLaris Kefersteinia orbieuLaris Kefersteinia parviLabris Kefe rste inia p ellita Kefersteinia retanae Kefersteinia taurina Kefersteinia taurina Kefersteinia toLùnensis Kefe rsteinia toLimensis Kefersteinia tolimensis Kefersteinia werekLei Koellensteinia g raminea LeoehiLus tricuspidatus LeucohyLe subuLata Loekhartia amoena Loekhartia pittieri Lyeasle Lyeaste Lyeaste Lycaste Lyeaste Lyeaste Lyeaste deppei P0230 Lycaste deppei P0460 P0437 P0024 P0334 P0476 P0201 P0383 P0391 P0129 P0180 P0357 P0361 P0364 P0235 P0178 P0240 P0028 PO 107 PO 103 POIOI P0167 P0262 P0095 P0234 POI04 Lyeaste Leueantha Lycaste leucantha Lyeaste Leucantha Lycaste x Michelii Lyeaste sehilleriana Lycaste tricoLor Lyeaste tricoLor M asdevallia reiehenbachiana M axillaria Max illaria M axilla ria M axilla ria Maxillaria Max illa ria Max illaria M axillaria Maxillaria Max illaria aeostae Maxillaria aLba Maxillaria angustissima Maxillaria araehnitiflora Maxillaria araehnitifiora Maxillaria atwoodiana Max illaria atwoodiana M axillaria braehybulbon Country Costa Rica Costa Rica Philippines Costa Rica Ecuador Panama Costa Ri ca Costa Rica Costa Rica Ecuador Costa Rica Costa Rica Costa Rica Costa Rica Ecuador Costa Rica Ecuador Ecuador Colombia Colombia Colombia Costa Rica Ecuador Costa Rica Nicaragua Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica M éxicoNicarag ua MéxicoNicaragua Costa Rica Costa Rica Costa Rica Panama Costa Ri ca Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Collector Bogarfn 806 DressLer 6831 Bogarfn 761 Pupulin 5029 BLanco 1053 Pupulin 655/ Ho Lst 8773 Pupulin s.n. PupuLin s.n. Bogarfn 143 Pupulin 5998 Cambronero s.n. Pupulin 6550 Pupulin 5892 Pupulin 4205 Pupulin 4205 Pupulin 4205 PupuLin 3039 PupuLin 6517 PupuLin 5648 PupuLin 4301 PupuLin 4515 BLanco 2653 Pupulin 2780 Date 13.05.2005 19.07.2007 13 .05.2005 09.05.2006 02.05.2006 12.03.2006 19.07.2007 13.05.2005 13.05.2005 19.07.2007 07.03.2006 13.05.2005 13.05.2005 13.05.2005 02.05.2006 06.06.2006 19.07.2007 17.03.2006 30.10.2002 19.07 .2007 13.05.2005 13.05.2005 20.07.2007 07.07.2005 09.06.2005 13.05 .2005 13.05.2005 22. 12.2005 29.12.2005 03 .05.2006 07.03.2006 20.07.2007 20.07.2007 20.07 .2007 06.06.2006 Bogarfn 2070 Vega s.n. BLanco 2808 BLanco 2808 G6mez 40 Pupulin 3023 Whitten 2045 BLanco 941 PupuLin 2850 Bogarfn 344 BLanco 2262 Bogarfn 874 20.07.2007 20.07 .2007 03.10.2002 26.10.2006 19.08.2007 09.05 .2006 07.03.2006 07.03 .2006 22.1 2.2005 22. 12.2005 14.11.2006 14.11.2006 14.1 1.2006 06.06.2006 22.1 2.2005 13.06.2006 05 .05.2006 19.05.2005 19.05.2005 19.05.2005 22.12.2005 30.06.2006 18.05.2005 06.06.2006 19.05.2005 76 TABLE No. P0249 P0189 P0268 PO 179 P0259 P0236 P0365 P0181 P0097 P0264 P0093 P0176 P0106 POO09 P0244 P0099 P0248 PO 102 PO 108 POI77 P0094 P0242 POO96 POOlO P0366 POlOO P0098 P0232 P0239 PO 109 POl57 P0321 P024 1 P0243 P0371 P0369 P0370 P0470 P0074 P0080 P0078 P0079 POO77 P0382 P0443 P0223 P0023 P0190 P0081 P0215 P0214 P0034 P0375 P0434 P0373 P0372 P0374 P0432 P0431 P0331 P0182 P01I3 SELBYANA 1. Volume 29(1) 2008 Continued. Genu s/species Maxillaria brunnea Maxillaria caespitifica Maxillaria campanulata Maxillaria campanulata Maxillaria confusa Maxillaria con/usa Maxillaria con/usa Maxillaria endresii Maxillaria endresii Maxillaria haberi Maxillaria hedwigae Maxillaria inaudita Maxillaria lueri Maxillaria luteoalba Maxillaria meridensis Maxillaria moralesii Maxillaria moralesii Maxillaria nasuta Maxillaria pseudoneglecta Maxillaria punctostriata Maxillaria ramonensis Max!llaria reichenheimiana Maxillaria reichenheimiana Maxillaria ringens Maxillaria ringens Maxillaria ringens Maxillaria rodrigueziana Maxillaria rodrigueziana Maxillaria sanguinea Maxillaria schlechteriana Maxillaria scorpioidea Maxillaria serrulata Maxillaria tonduzii Maxillaria wercklei Mesospinidium horichii Mesospinidium panamense Mesospinidium warszewiczii Micochilus bimentatus Miltonia regnellii Miltoniopsis phalaenopsis Miltoniopsis roezlii Miltoniopsis vexillaria Miltoniopsis warscewiczii Mormodes colossa Mormodes fractiflexa Mormodes horichii Mormodes atropurpurea Mormodes colossa Mormolyca ringens Oncidium Oncidium Oncidium coquianum Oncidium dichromaticum Oncidium dichromaticum Oncidium ensatum Oncidium fuscatum Oncidium hyphaematicum Onc idium hyphaematicum Oncidium kotzchianum Oncidium luteum Oncidium luteum Oncidium oliganthum Country Costa Rica Costa Rica Costa Ri ca Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Colombia Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Panama Costa Rica Costa Rica Brazil Colombia Colombia Colombia Costa Rica Costa Rica Costa Rica Costa Rica. Costa Rica Costa Rica Costa Rica Costa. Rica Costa Ri ca Costa Rica Costa Rica Costa. Rica Ecua.dor Nicaragua Nicaragua N icaragua. Costa Rica Costa. Rica. Costa Rica Guatemala Collector Whitten 2028 Blanco 1076 Bogarfn 1801 Comez 48 Bogarfn 1324 Whitten 2030 Whitten 2030 Whitten 2089 Bogar{n 1868 Pupulin J 859 Comez S.n. Comez 19 Pupulin 4682 Pupulin 3247 Comez 1 Blanco 2107 Comez 24 Comez 24 Whitten 2/81 Bogarfn 1135 Blanco 3179 Blanco 3179 Pupulin 5357 Dressler s.n. Pupulin 6690 Blanco 2691 Bogarfn 2705 Bogarfn 2561 Blanco 2192 Comez 18 Pupulin 3438 Pupulin 2495 Pupulin 6421 Pupulin 6420 Pupulin 6422 Bogarfn 3793 Pupulin 5455 Pupulin 6197 Whitten 2192 Pupulin 3378 Date 16.06.2006 22.12.2005 12.07.2006 22.12.2005 30.06.2006 06.06.2006 14.1 1.2006 22.12.2005 18.05.2005 05.07.2006 18.05.2005 22.12.2005 19.05.2005 06.11.2002 13.06.2006 18.05.2005 16.06.2006 19.05.2005 19.05.2005 22.12.2005 18.05.2005 13.06.2006 18.05.2005 21.1 1.2002 14.11.2006 18.05.2005 18.05.2005 06.06.2006 07.06.2006 19.05.2005 22.12.2005 03.10.2006 13.06.2006 13.06.2006 25.01.2006 25.01.2006 25.01.2006 13.08.2007 13.05.2005 13.05.2005 13.05.2005 13.05.2005 13.05.2005 07.03.2006 20.07.2007 12.05.2006 04.03.2005 22.12.2005 13.05.2005 03.05.2006 03.05.2006 19.04.2006 28.02.2006 19.07.2007 05.02.2006 05.02.2006 05.02.2006 19.07.2007 19.07.2007 26.10.2006 22.12.2005 24.05.2003 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER T AB LE No. POIIl P0458 POl83 P0440 P0418 P0333 POI14 P0250 P0266 P0082 P0273 P0455 POl61 P0446 P0389 P0068 P0061 POO07 P0062 POI92 POO02 P0173 P0397 POO18 P0472 P0022 P0265 P0314 P0063 P0392 POl34 POl62 P0133 P0116 P0158 P0270 P0083 POll2 P0135 POOll P0463 P0445 P0136 P0209 P0433 P0438 P0233 P0356 P0338 P0306 P0337 P0363 P0260 P0367 P025 1 P0492 P0343 P0259 P0256 P0160 P0448 P0263 1. 77 Continued. Genus/species Oncidium parviflorum Oncidium parviflorum Oncidium schroederianum Oncidium schroederianum Oncidium sp.nov. Oncidium stenobulbon Oncidium storkii Osmoglossum egertonii Otoglossum chiriquense Otoglossum globuliferum Paphinia subclausa Paphinia subclausa Paphinia subclausa Pescatorea cerina Pescatorea cerina Pescatorea cerina Pescatorea cerina Pescatorea cerina Pescatorea dayana Phalaenopsis amabilis Phalaenopsis schilleriana Phalaenop sis amabilis Phalaenopsis luedemanniana Phalaenopsis violacea Polycicnis barbata Polycycnis barbata Polycycnis barbata Polycycnis barbata Prosthechea brassavolae Prosthechea chacaoensis Prosthechea neglecta Prosthechea chacaoensis Prosthechea vitellina Rodriguezia venusta Rodriguezia venusta Rossioglossum schlieperianum Rossioglossum schlieperianum Rudolfiella aurantiaca Schomburgkia lueddemannii Sedirea japonica Sievekingia fimbriata Sievekingia fimbriata S ieveking ia fimbriata Sievekingia fimbriata Sievekingia suavis Sigmatostalix cardioglossa Sigmatostallx dulcineae Sobralia Sobralia Sobralia Sobralia Sobralia Sobralia amabilis Sobralia andreae Sobralia hel/eri Sobralia luteo la Sobralia macra Sobralia macrophyl/a Sobralia quinata Sobralia leuçoxantha Stanhopea Stanhopea Country Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rfca Costa Rica Costa Rica Costa Rica Costa Rica Colo mbi a Asia Philippines lndonesia Philippines Malasya Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Mexico Ecuador Ec uador Costa Rica Costa Rica Ecuador Costa Rica Japan Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa, Rica Costa Rica Costa Rica Colombi a Costa Rica Costa Rica Costa, Rica Costa Rica Costa, Rica Costa Rica Costa Rica Costa Rica Collector Bogarfn 2468 Pupulin 6254 Karremans 1650 Bogar(n 1065 Pupulin 6235 Pupulin 5296 Dressler 6590 Dressler 6590 Pupulin 4665 Blanco 2642 Pupulin 2993 Pupulin 2097 Pupulin 5713 Pupulin 5713 Bogarfn 1898 Bogarfn 1898 Bogarfn 1845 Pupulin 5281 Bogarfn 2851 Bogarfn 2813 Bogarfn 2575 Pupulin 3499 Bogar(n 2079. Aguilar 13 Bogarfn 913 Dressler 6723 Dressler 6621 Karremans 661 Dressler 671 3 Dressler 6539 PupiAlin 2629 Casteifranco 155 Pupulin 3664 Pupulin 4184 Date 24.05 .2003 20.07 .2007 22.1 2.2005 20.07.2007 19.07 .2007 26.10.2006 24.05.2003 16.06.2006 12.07.2006 13.05.2005 01.09.2006 20.07.2007 07.07.2005 20.07.2007 07.03.2006 17.07 .2005 13.05.2005 06.08.2003 13.05.2005 17.03 .2006 12.05 .2004 17.07.2005 28.03.2006 22.10.2005 19.08.2007 04.03.2005 12.07.2006 29.09.2006 22.12.2005 07.03 .2006 10.06.2005 07 .07.2005 10.06.2005 08.06.2005 16.05 .2005 18.07.2006 13 .05.2005 24.05 .2003 13.06.2005 12 .05 .2004 20.07.2007 20.07.2007 13.06.2005 03.05.2006 19.07.2007 20.07.2007 06.06.2006 14.11.2006 14 .11.2006 13.09.2006 14.11.2006 14.11.2006 30.06.2006 14.11.2006 21.06.2006 05 .02.2008 14.11.2006 30.06.2006 30.06.2006 07.07.2005 20.07 .2007 05.07.2006 TABLE,1. Continued. Genus/species No. P0267 P0294 P0430 P0452 P0277 P0387 P0276 P0291 P0280 P0285 P0292 P0210 P0257 P0247 P0290 P0204 P0203 P0295 P0315 P0485 P0466 P0225 P045 1 P0457 P0474 P0205 P0368 P0254 P0283 P0288 P0289 P0137 P0305 P0471 P0202 P0253 P0252 P0459 P01 40 P0449 P0330 P0345 P0353 P0206 P0208 P0020 P0344 P0328 P0335 P0296 P0484 POO16 P0146 P0 147 P0207 P0482 P0483 P0487 POO19 P0138 P0139 P0 14 1 P01 42 Volume 29(1) 2008 SELBYANA 78 Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stmihopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea Stanhopea cfcostaricensis cirrhata cirrhata cirrhata cirrhata cirrhata confusa costaricensis costaricensis costaricensis costaricensis costaricensis costaricensis costaricensis x cirrhata ecornuta ecornuta ecornuta ecornuta ecornuta ecornuta ecornuta x fowlieana x fowlieana gibbosa gibbosa gibbosa grandiflora oculata oculata pulla pulla pulla pulla wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii wardii Country Costa Rica Costa Rica Costa Rica Costa Rica Costa R ica Costa Rica Costa Rica Costa Rica Costa Rica Ecuador Nicaragua Venezuel a Venezuela Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica C osta Rica Costa Rica Costa R ica Costa Rica Costa Rica Costa Rica Costa R ica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Mexico M exico Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa R ica Costa Rica Collector Bogarfn 1639 Bogarfn 757 Pupulin 2960 Pupulin 2960 R ojas 112 Pupulin 6279 Karremans 557 Bogarfn 1021 Bogarfn 1021 Castelfranco 153 Pupulin 2843 P up ulin 5827 Flores S.n. Rojas 113 Rojas 110 Rojas 134 Rojas 135 Roj as 136 Bogarfn 630 Dressler 6564 G6mez 51 Pupulin 4930 Pupulin 4390 Pupulin 11 79 Pupulin 4930 P upulin 1179 Date 12.07.2006 05.09.2006 19.07.2007 20.07.2007 01.09.2006 07 .03.2006 01.09.2006 05 .09.2006 05.09.2006 05.09.2006 05.09.2006 03.05 .2006 30.06.2006 13.06.2006 05.09.2006 03.05.2006 03.05.2006 05.09.2006 29 .09.2006 02.10.2007 20.07.2007 16.05.2006 20.07.2007 20.07.2007 19.08.2007 03.05 .2006 26.11.2006 2 1.06.2006 05 .09.2006 05.09.2006 05.09.2006 13.06.2005 13.09.2006 19.08.2007 03 .05.2006 21.06.2006 21.06.2006 20.07.2007 13.06.2005 20.07.2007 26.10.2006 14.11.2006 14. 11.2006 03.05.2006 03.05 .2006 21.06.2003 14.11.2006 17.10.2006 26.10.2006 05 .09.2006 02.10.2007 20.07.2004 13.06.2005 13.06.2005 03.05.2006 28 .09.2007 28.09.2007 25.10.2007 05 .08.2003 13.06.2005 13.06.2005 13.06.2005 13.06.2005 79 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER TABLE No. P0144 P0145 P0293 P0297 P0304 P0327 P0143 P0269 P0348 P0350 P0279 P0318 P0415 P0191 P0407 P0199 POl52 P0066 P0065 P0421 P0245 P0477 P0064 P0422 P0423 P0384 P0435 P0298 P0237 P0238 P0148 P0073 P0070 P0220 P0323 P0211 P0085 P0084 P0377 P0360 P0424 P0349 P0490 P0399 P0491 P0486 P0351 POl69 P0400 POl51 P0076 P0175 P0309 P0453 P0075 poon P0320 P021 2 P0213 P04 4 P0059 P0058 P0401 1. Continued. Genus/species Stanhopea wardii Stanhopea wardii Stanhopea warscewicziana Stanhopea warscewicziana Stanhopea warscewicziana Stanhopea warscewicziana Stanhopea warscewicziana Stenia Stenia calceolaris Stenia dodsoniana Stenia guttata Stenia pallida Stenia pallida Stenia stenioides Ste nia stenioides Stenotyla lankesteriana Stenotyla lankesteriana Stenotyla lendyana Stenotyla lendyana Stenoty la lendyana Stenotyla panamensis Stenoty la panamensis Stenotyla picta Stenoty la picta Stenotyla picta Stenotyla picta Stenotyla picta TeLipogon biolleyi Telipogon costaricensis Telipogon monticola TeLipogon ampliflorus Ticoglossum krameri Ticoglossum oerstedii Tolumnia triquetra Trevoria zalhbrukneriana Trevoria zalhbrukneriana Trichocentrum capistratum Trichocentrum cymbiglossum Trichocentrum pfavii Trichoglottis Trichopilia TrichopiLia Trichopilia Trichopilia eneidae Trichopilia fragrans Trichopilia fragrans Trichopilia aff. fra grans Trichopilia galeottiana Trichopilia hennisiana Trichopilia marg inata Trichopilia marg inata Trichopilia marginata Trichopilia nobilis Trichopilia olmosii Trichopilia punicea Trichopilia punicea Trichopilia turrialbae Vanda coerulea Vanda lamellata Warcze wiczella discolor Warsce wiczella discolor Warcze wiczella ionoleuca Warczewiczella ionoleuca Country Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Ecuador Ecuador Ecuador Ecuador Ecuador Ecuador Ecuador Ecuador Costa Rica Costa Rica Guatemala Guatemala Guatemala Panama Panama Costa Rica Costa Rica Costa Rica Costa Rica Panama Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Costa Rica Jamaica Costa Rica Costa Rica Costa Rica Costa Rica Costa Ri ca Philippines Colombia Colombia Costa Rica Panama Colombia Colombia Colombia Costa Rica Colombia Costa Rica Costa Rica Costa Rica Colombia Panama Costa Rica Costa Rica Costa Rica Philippines Philippines Costa Rica Costa Rica Ecuador Ecuador Collector Pupulin Pupulin Pupulin Pupulin Pupulin 6239 6332 6340 5937 6286 Pupulin 5893 Pupulin 5893 Hubein s.n. Pupulin 5868 Pupulin 5869 Pupulin 3031 Dressler s.n. Bogarfn 2960 Karremans 13 17 Rojas 7148 Rojas 7157 Bogarfn 1900 G6mez 39 Pupulin 2075 PupuLin 4754 Pupulin 5826 Dressler 6807 PupuLin 6333 Pupulin 641 8 Pupulin 6341 Pupulin 6341 Bogarfn 145 Pupulin 4382 Pupulin 4433 Bogarfn 2948 Pupulin 5213 PupuLin 5235 PupuLin 5235 Dale 13.06.2005 13.06.2005 05.09.2006 06.09.2006 13.09.2006 10.10.2006 13.06.2005 18.07.2006 14.1 1.2006 14.11.2006 05.09.2006 29.09.2006 19.07 .2007 17.03.2006 19.07.2007 02.05 .2006 19.04.2006 13.05.2005 13.05.2005 19.07.2007 13.06.2006 19.08.2007 13.05.2005 19.07.2007 19.07.2007 07.03.2006 19.07.2007 06.09.2006 06.06.2006 06.06.2006 13.06.2005 13.05.2005 13.05.2005 03.05.2006 03 .10.2006 03.05.2006 13.05.2005 13.05 .2005 05.03.2006 14.11.2006 19.07.2007 14.11.2006 14.01.2008 28.03.2006 14.01 .2008 25 .10.2007 14.11 .2006 07.07 .2005 28.03 .2006 14.06.2005 13.05 .2005 17.07.2005 20.09.2006 20.07 .2007 13.05 .2005 13.05.2005 03. l0.2006 03 .05 .2006 03.05.2006 19.07.2007 13.05.2005 13.05.2005 19.07.2007 80 SELBYANA Volume 29(1) 2008 Record: 470 Total: 493 Unsort ed RASTREO ( SCAt> IONUf,I,srn GENUS P0470 JBL-INVENT JBL-s .n . Mfcrochilus FAMI:..Y '5PECIES Orchidaceae bimentotus INflIASP COUtfTRY Costa Rica . COLlDATE 30 .07 .2007. COLLECTOR F. Pupulln COLLNUH.8ER 6690. TAXONOfl Det . F. Pupulin, 8.2007. DATADATE POll.MAT JBL-PoIIColl. HERSM.\T FAMAT JBl-Splrlt. 13.08.2007. ILWSTR.'AAT Ｑｉｩ｜ｬｾｲ､｡＠､ｾ＠ Co ... ta Rica .., Record: Total: 493 Un50rted RASTREO (S(AN IDNUI4ER GENUS P0492 JBL- 09442 ｪｂｌﾷｉｾｎｅｔ＠ Sabra/ia FAMlLY SPECIES Orchidaceae /uteolo INfRASP (QUtfTRY Costa Rica . 10.12.2000. COlLECTOR: F. Pu pulin TAXONOE'T Det. R. l. Dressler,03.2007. POLlMAT JBl-polleaU. HERaMAT FIGURE 4. COlLNIJ.'.\8E:R 2629 . DATADATE 05 .2.2008 . 'MAAT ILlUSTRMAT Foto. Two pages from the pollinaria database at Lankester Botanical G arden. PUPULIN & KARREMANS: POLLINARIA AT LANKESTER .......·ea.... ..... ﾷｾ＠ ..... - ......... ($! .......... _ · ... *' ... _,-. ... _,..... 81 !_·. __;___ . _C>Ol".·"'_"'" ........ • • • • • 11 • • • FIGURE 5. Two pages from "EPIDENDRA," the online taxonomie and nomenc\atural database of Lankester Botanical Garden. mize the damage to the viscidium or the caudicles. This is to permit the documentation of the pollinaria from as many views as possible. After removal, the pollinaria are scanned on a flatbed scanner (Epson Perfection 4490 Photo and Epson Perfection V200 Photo) at 2400 to 3200 dpi resolution, and saved as TIFF files including relative scales. Dorsal and ventral views are recorded (FIGURE 1), and a lateral view is included when feasible (FIGURE 2). The pollinaria are then stored in specifically designed white paper envelopes and kept in hermetic plastic boxes (FIGURE 3). The internai humidity of the boxes is controlled by means of silica gel envelopes. High-resolution digital images of the pollinaria (approx. 2 Mb) are permanently stored in exter" nal hard drives, while low-resolution images (approx. 170 Kb) are included as visual references in a database designed for that purpose (FileMaker Pro 9.0) (FIGURE 4). This includes locality and collecting data, as weil as a crossreference to the spirit-preserved vouchers from which pollinaria were removed. Copies of the entire database on CD may be sent on request to researchers and institutions, addressing the correspondence to the senior author. In order to improve public access to the information of the database, digital images of selected pollinaria from the collection are progressively displayed in the main taxonomic and nomenclatorial database of LBG, "EPIDENDRA," which will be available soon through the Internet (FIGURE 5). The collection at LBG actually includes 496 pollinaria, belonging to 312 species in 94 genera (Table. 1). Although the collection also includes pollinaria of the subfamily Orchidoideae (subtribes Cranichidinae and Spiranthinae), the compld pollinaria of the Epidendroideae are the best represented. In this subfarnily, we have representative species of the Sobralieae (13 pollinaria), Calypsoeae (1), Epidendreae (23), Cymbidieae (427), and Vandeae (19) (TABLE 2). Par- ticularly weil represented are pollinaria from the more highJy evolved Neotropical groups, where the pollinia are accompanied by distinct viscidia and stipes, with emphasis on the subtribes Oncidiinae (89 pollinaria), Maxillarinae (69) , Stanhopeinae (110), and Zygopetalinae (149). Also present are pollinaria of Paleotropical Epidendroideae, among which are species of Aeridinae (16), Angraecinae (2), Aerangidinae (1), and Dendrobiinae (3) . Works are in progress at LBG to use the large collections of pollinaria of the Zygopetalinae (FIGURE 6) to reconstruct phylogenetic tendencies in the organization of pollen and associate structures in this subtribe (Pupulin in prep.), and designed to use pollinaria morphology for species identification in the genus Stanhopea (FIGURE 7). ACKNOWLEDGMENTS We are indebted to all the colleagues at LBG who helped through the years documenting pollinaria for collection, and in particular to Diego Bogarfn, Gustavo Rojas, Hilda Le6n-Paez, Jaime Aguilar, and Emily Serrano. We thank Margherita Pupulin and Carlotta Pupulin, who devoted part of their summer holidays to substantially improve the collection. All the pollinaria included in the collection were removed from plants collected in the field or obtained through scientific interchange, thanks to scientific passports and the import permits issued by the Ministry of the Environment of Costa Rica (MINAE) and its National System of Protected Areas (SINAC). We sincerely acknowledge their continuous cooperation. The Orchid Pollinaria Collection at LBG is part of the general Collections of the University of Costa Rica, and its maintenance and management are supported by the University Collections Committee. 82 SELBYANA Volume 29(1) 2008 F IGURE 6. Representative pollinaria of the Zygopetalinae. 1-8: Dichaea. 1-D. truila (Pupulin 5130); 2D. calyculata (Pupulin 6923); 3-D. globosa (Pupulin 5319); 4-D. globosa (JBL-09172 ); 5-Dichaea sp. (Pupulin 6891) ; 6-D. morrisii (Whitten 2171); 7 - D. poiciilantha (Karremans 1140) ; 8- D. obovatipetala (JBL-08203) . 9-13: Daiotyla. 9-D. xanthina (Pupulin 5891); lü,lI- D. albicans (Pupulin 4372); 12, 13-D. crassa (Pupulin 4257). 14- 16: Chondroscaphe. 14-C. amabilis (Pupulin 6553); 15- D. escobariana (Pupulin 6289); 16-D. flav eola (Pupulin 6474). 17-27: Kefersteinia. 17- K. costaricensis (Bogarfn 761); 18-K. laminata (Pupulin 6551); 19- K. o rbicularis (Pupulin 2068); 20-K. retanae (JBL-011 13); PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 83 (Continued) 2 1-K. wercklei (JBL-03465); 22- K. excentrica (Pupulin 5225); 23-K. excentrica (JBL-00469); 24-K. parvilabris (JBL-05186); 25-K. tolimensis (Pupulin 4205); 26-K. mystacina (Pupulin 6559); 27- K. taurina (Pupulin 6550). 28-32: Benzingia. 28- B. reichenbachiana (Bogarîn 1890); 29, 30-B. reichenbachiana (JBL-07994); 31-B. cornuta (Pupulin 5879); 32- B. caudata (Pupulin 6544). 33: xBensteinia ramonensis (Bogarîn 1923). 34: Batemannia leferenzii (Pupulin 6224). 35: Zygopetalinae gen. ined. (Pupulin 6605). 3641 : Chauba rdiella. 36-C pubescens (Pupulin 6240); 37-C aff. pubescens (Pupulin 6566); 38-C subquadrata (Pupulin 5716); 39, 40-C pacuarensis (JBL-0111 7, JBL-05422); 4t-C aff. subquadrata (Pupulin 6564). 42- 48: Stenotyla. 42, 43- S. lankesteriana (JBL-OJ651); 44, 45-S. picta (JBL-OJ970, JBL-01905) ; 46, 47- S. lendyana (JBL-02053, JBL-0896); 48-S. panamensis (Pupulin 5868). 49- 56: Stenia. 49- S. guttata (Pupulin 6567); 50-S. dodsoniana (Pupulin 6340) ; 51 -S. stenioides (Pupulin 5893); 52-S. angustilabia (Pupulin 6570) ; 53-S. calceolaris (Pupulin 6332); 54, 55-S. pallida (Pupulin 6286; Pupulin 6548); 56-S. lillianae (Pupulin 6557). 57-62: Warczewiczella. 57, 58- W. discolor (Pupulin 3939; JBL-04925); 59- W. marginata (Pupulin 6516); 60, 61-W. ionoleuca (Pupulin 5235; JBL-00873);62- W. palatina (Pupulin 6909). 63-65 : Cochleanthes. 63, 64-C aromatica (JBL-02932; JBL-07583); 65-C fla belliformis (Pupulin 6621). 66: Zygosepalum lindeniae (Pupu lin 6643). 67: Ixyophora viridisepala (Pupulin 6554). 68: "Chondrorhycnha " velastiguii (Pupulin 6622). 69- 72: Pescatorea. 69, 70, 71 - P. cerina (JBL-09435; JBL-09430 ; JBL-06217); 72- P. dayana (JBL-03943). 73: Zygopetalum intermedium (Pupulin s.n.). 74: Galeottia grandiflora (JBL-J4627). 7578: Huntleya. 75, 76-H. burtii (JBL-14492; Pupulin 3829); 77-H. citrina (Pupulin 6645); 78-H. lucida (Pupulin 6642). 79-80: Euryblema andreae (Pupulin 6282). Scale bar = 5 mm. 84 SELBYANA Volume 29(1) 2008 FIGURE 7. Pollinaria of Stanhopea spec ies. 1- 16: S. wardii. I- Dressler 6564; 2 - JBL-04360; 3 - JBL06666; 4 - DB 630 ; 5 - JBL-07741; 6 - R . Gomez 51; 7 - JBL-06133 ; 8 - FP 1179; 9 - JBL-04363; 10 - FP 4390; 11 - JBL-09981 ; 12 - JBL-07743 ; 13 - FP 4930; 14 - JBL-06126; 15 - JBL-0615J ; 16 - FP 4930. 1721: S. warscewicziana.17 - JBL-14610; 18 - JBL-08955; 19 - JBL-06148; 20 - JBL-14605 ; 21 - JBL-07747. 2228: S. costaricensis. 22 - JBL-05094 ; 23 - JBL-05659 ; 24 - JBL-07726; 25 - JBL-05094 ; 26 - JBL-05066; 27 JBL-09988; 28 - JBL-05069. 29-35: S. ecomuta. 29 - JBL-09867; 30 - JBL-1l 353; 31 - JBL-12602; 32 - JBLJJ696; 33 - JBL-06132 ; 34 - Rojas 110; 35 - JBL-11698. 36- 39: S. pulla. 36 - JBL-05657; 37 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 2 . Po lli naria included in the coll ection, arranged systematically (phylogentic c lassification according to Chase et al. 2003). Numbers in brackets refer to the number of pollinaria for each spec ies or morphospecies. TABLE Orchidoideae Cranichideae Goodyerinae: Erythrodes tridax, Microchilus bimentatus Spiranthinae: Coccineorchis standleyi Epidendroideae Sobralieae: Sobralia amabilis, S. andreae, S. helleri, S. luteola, S. mac ra, S. macrophylla, S. quinata, S. leucoxantha, Sobralia ssp. (5) Calypsoeae: Govenia ciliilabia Epidendreae Pleurothallidinae: Masdevallia reichenbachiana Laeliinae : Arpophyllum giganteum, Barke ria lindleyana, Brassavola nodosa, Cattleya inlermedia, Encyclia ambigua, E. cordigera, E. ceratistes, Epidendrum centradenia, E. odontochilum, E. schumannianum, E. wallisii, Epidendrum spp. (3), Guarianthe skinneri, Prosthechea brassavolae, P. chacaoensis (2), P. neglecta, P. vitellina, Schomburg kia lueddemannii Coeliinae [?): Coelia bella Cymbidieae Catasetinae: Catasetum maculatum (3), Cycnoches amparoanum, Cyrtopodium macrobulbon, Dressleria dilecta, Galeandra lacustris, Mormodes atropurpurea, M. colossa (2), M. fra ctiflexa, M. horichii Cymbidiinae: Cymbidium iridioides (2), Grammatophyllum multiflorum, G. scriptum Eriops idin ae: Eriopsis wercklei (2) Oncidiinae: Ada aurantiaca (2), A. chlorops, Aspasia epidendroides, Brassia arcuigera (6), B. caudata (4), B. gireoudiana (2), B. suavissima, B. verrucosa, Comparettia falcata, Erycina cristagalli, Leochilus tricuspidatus, Lockhartia amoena, L. pittieri, Mesospinidium horichii, M. panamense, M. warszewiczii, Miltonia regnellii, Miltoniopsis phalaenopsis, M. roezlii, M. vexillaria, M. warscewiczii, Oncidium coquianum, O. dichromaticum, O. dichromaticwn, 0. ensatum, O. fuscatum, O. hyphaematicum (2), 0. klotzchianum, O. luteum (2), O. oliganthum, O. parviflorum (2), O. parviflorum, O. schroederianum (2), 0. stenobulbon, O. storkii, Oncidium spp. (3), Osnwglossum egertonii, Otoglossum chiriquense, 0. globuliferum, Rodriguezia venusta (2), Rossioglossum schlieperianum (2), Sigmatostalix dulcineae, S. cardioglossa, Telipogon ampliflorum, T. biolleyi, T. costaricensis, T. monticola, Ticoglossum krameri, T. oerstedii, Tolumnia triquelra, Trichocenlrum capistratum, T. cymbiglossum, T. pfavii, TABLE 2. 85 Continued. Trichopilia eneidae, T. fragans (2), T. aff. fragrans, T. galeottiana, T. hennisiana, T. marginata (3), T. nobilis. T. olmosii, T. punicea (2), T. subuLata, T. turrialbae, Trichopilia spp. (3) Maxillariinae: Helerotaxis sessilis, Lycasle borucana (2), L. depp ei (2), L. leucantha (3), L. michelii (lassioglossa X cochleata), L. schilleriana, L. tricolo r (2), Lycaste spp. (4), Maxillaria acostae, M . alba, M. angustissima, M . arachnitiflora (2) , M. atwoodiana (2), M. brachybulbon, M. cf. brunnea, M. caespitifica, M. campanulata (2), M. confuse (3), M. endresii (2) , M. haberi, M. hedwigae, M. inaudita, M. lue ri, M. luteoalba, M. meridensis, M. moralesii (2) , M. nasuta, M. pseudoneglecta, M. punctostriata, M . ramonensis, M. reichenheimiana (2), M. ringens (3), M. rodrigueziana (2), M. sanguinea, M. schlechteriana, M . scorpioidea, M. serrulata, M. tonduzii, M. wercklei, Maxillaria spp. (9), Mormolyca ringens, Rudoljielia aurantiaca Stanhopeinae: Acineta densa (7), Acineta sp. (2) , Gongora armeniaca (2), G. claviodora, G. galeata (2), G. leucochila (2), G. truncata, G. unicolor (2), Gongora spp. (6), Houlletia tigrina, Paphinia subclausa (3), Polyc icnis barbata (4), Sieveking iajimbriata (3), S. sua vis, Stanhopea cirrhata (5), S. confusa, S. costaricensis (6), S. cf. costaricensis, S. costaricensis X cirrhata, S. ecornuta (7) , S. gibbosa (3), S. grandiflora, S. oculata (2), S. pulia (4), S. wardii (19) , S. warscewicziana (5) , S. X fowlieana (2), Stanhopea spp. (14), Trevoria zalhbrukneriana (2) Coeliopsidinae: Coeliopsis hyacinlhosma (2) Zygopetalin ae: Batemania leferenzii, X Bensteinia ramonensis (2) , Benzing ia caudata, B. cornuta, B. reichenbachiana (2), Chaubardia heteroclita, Chaubardiella pubescens, C. subquadrata, C. pacuarensis (2), Chondroryncha thienii (2) , Chondroscaphe amabilis, C. atrilinguis (2), C. bicolor (9), C. aff. bicolor (15), C. endresii (2), C. eburnea (2), C. embreei (2), C. escobariana, C. flaveola (3), C. cf. laevis (2), C. yamilethiae (5), Cochleanthes aromatica (2), Daiotyla albicans (2), D. crassa (3), D. xanthina, Dichaea acroblephara (3), D. amparoana, D . cryptarrhena, D. elliptica (3), D. fragrantissima subsp. eburnea (2), D. globosa (3), D. morrisii (2), D. ovobatipetala (2) , D. panam ensis (2), D. poicillantha (3), D. similis, D. cf. similis, D. trulla (2), Euryblema anatonum, E. andreae (2), Galeottia grandiflora (3), Huntleya burtii (2) , Kefersteinia andreettae, K. auriculata, K. costaricensis (2), K. excentrica, K. laminata, K. orbicuLaris (2) , K. pan1ilabris, K. pellita, K. retanae, K. taurina(2), K. tolimensis (3), K. wercklei, Kefersteinia spp. (2), Koellensteinia g raminea, Pescatorea cerina (5), P. dayana, Stenic calceolaris, S. dodsoniana, S. guttata, Stenia pallida (2), Stenia stenoides (2), Stenotyla lankeste riana (2), S. len- f-- (Continued) - JBL-06155; 38 - Rojas 136; 39 - J BL-07727. 40-44: S. c irrhata. 40 - DB 1021; 41- DB 1021; 42 - FP 2843; 43 - FP 583; 44 - Castelfranco 153. 45- 47: S. gib bosa. 45 - JBL-04956; 46 - JBL-0 7529; 47 JBL-07529. 48 - S. confusa (JBL-09484). 49 - S. grandiflora (J BL-09988). 50, 51 - S. oculata (Rojas 134; Rojas 135). 52, 53 - S. x fowliana (JBL-1303l; JBL-1303l). 54 - S. costaricensis x cirrhata (Rojas 113). 55- 65: Stanhopea spp. 55 - JBL-05654; 56 - FP. 2960; 57 - JBL-09988 ; 58 - FP. 2960; 59 - AK 557; 60 - FP 6279; 61 - JBL-08970; 62 - JBL-l2004 ; 63 - JBL-08970; 64 - DB 1639; 65 - Rojas 112. Scale bar = 5 mm. 86 TABLE 2. SELBYANA Continued. dyana (3), S. panamensis, S. picta (5), Warczewiczella discolor (2), W. ionoleuca (2) Vandeae Aeridinae: Aerides falcata, A . multiflora, Asco centrum curvifolium, Gastrochilus calceolaris, G. formosanum, Haraella retrocalla, Hygrochilus parishii, Phalaenopsis amabi[js (2), P. lueddemanniana, P. schilleriana, P. violacea, Sedirea japonica, Trichoglottis sp., Vanda coerulea, V. lamellata Angraecinae: Angraecum ebumeum, A. eichlerianum Aerangiclinae: Eurychone rothschildiana Unplaced subtribes within Epidendroideae Dendrobiinae: Bulbophyllum lobbii, D endrobium antennuatum, Epigeneium cymbidioides LITERA TURE CITED Atwood , J.T. 1986. The size of the Orchidaceae and the systematic distribution of epiphytic orchids. Selbyana 9: 171- 186. Bogarfn, D. 2007. A new Lycaste (Orchidaceae: Maxillariinae) from Costa Rica. Lankesteriana 7(3): 543- 549. Borba, E .L. and J. Semir. 1999. Tçmporal variation in pollinarium size after its removal in species of Bulbophy llum: a different mechanism preventing self-pollination in Orchidaceae. Plant Syst. Evo\. 217 : 197-204. Chase, M.W. 1987. Systematic implications of pollinarium morphology in Oncidium Sw., Odontoglossum Kunth, and allied genera (Orchidaceae) . Lindleyana 2: 8-28. Chase, M .W. , KM. Cameron, R.L. Barrett, and J.v. Freudenstien. 2003. DNA data and Orchidaceae systematics: a new phylogenetic classification. Pp. 69-89 in KW. Dixon, S.P. Kell , R L. Barrett, and P.J. Cribb, eds. Orchid conservation. Natural History Publications, Kota Kinabalu, Sabah. Cribb, P. and R . Govaerts. 2005. Just how many orchids are there? Pp. 161- 172 in A . RaynalRoques, A. Roguenant, and D. Prat, eds. Proc. 18 World Orch. Conf. Naturalia Publ. , Turriers, France. Darwin, C. 1877. The various contrivances by which orchids are fertlized by insects. 2nd ed. John Murray, London. Dressler, R.L. 1968. Pollination by eug lossine bees. Evolution 22: 202- 210. - - - . 1976. How to study orchid pollination without any orchids. Pp. 534- 537 in K Senghas, ed. Proc. 8'" World Orch. Conf. ' Deutsche Orchideen Gesellschaft, Frankfurt. Volume 29(1) 2008 1981. The Orchids: Natural History and Classification. Harvard University Press, Cambridge, Mass. - - - . 1986. Features of pollinaria Wld orchid classification. Lindleyana 1: 125- 130. - - - . 1993. Phylogeny and ClassificatiQn of the Orchid Farnily. Timber Press, Portland. - - - and W. KJilcunas. 2006. A Sectional Classification of Trichopilia Lindley (Orchidaceae), Selbyana 27: 30- 33. Freudenstein, J.v. and F.N. Rasmussen. 1997. Sectile pollinia and relationships in the Orchidaceae PI. Syst. Evol. 205: 125- 146. Johnson, S.D. and L.A. Nilsson. 1999. Pollen carryover, geitonogamy and the evolution of deception in orchids. Ecology 80: 2607-2619. Johnson , S.D. and T.J. Edwards. 2000. The structure and fJ.]nction of orchid pollinaria. Plant Syst. Evo!' 222: 243-269. Nilsson, L.A. 1980. The pollination ecology of Dactylorhiza sambucina (Orch idaceae). Bot. Noti ser 133: 367-385. - - - . 1983. Processes of isolation and introgressive interplay between Platanthera bifolia (L.) Rich. and P. chlorantha (Custer) Reichb. (Orchidaceae). Bot. J . Linn. Soc. 87: 325- 350. Pupulin, F. 2007a. Contributions toward a reassessment of Costa Rican Zygopetalinae (Orchidaceae). A systematic revision of Dichaea in Costa Rica. Harvard Pap. Bot. 12(1): 15- 154. - - - . 2007b. Those violet things: the elusive Dichaeas of Costa Rica, an overview of Dichaea phylogeny and classification. 14'" EOCCE Proceedings. Caesiana n. S. 4: 37-49. - - - . In press. The natural and taxonomic history of Chondrorhyncha (Orchidaceae-Zgopetalinae). Proc. 19'" World Orch. Conf. Miami. Reichenbach, H .G . 1852. " De pollinis orchidearum genesi ac structura et de orchideis in arte ac systema ricligendis" Ph.D. diss. , Leipzig. Steiner, KE. 1989. The pollination of Disperis (Orchidaceae) by oil-collecting bees in southern Africa. Lindleyana 4:164- 183. Widmer, A. , S. Cozzolino, G . Pellegrino, M . Soliva, and A. Dafni. 2000. Molecular analysis of orchid pollinaria and pollinaria remains found on insects. Molec. Eco!' 9: 1911 - 1914. Williams, N.H. 1970a. Some observations on pollinaria in the Oncidiinae. Amer. Orch. Soc. Bull. 39: 32-43. - - - . 1970b. Sorne observations on pollinaria in the Oncidiinae- II. Amer. Orch. Soc. Bull. 39: 207220. - - - . 1972. Additional studies on pollinaria in the Oncidiinae. Amer. Orch. Soc. Bull. 41: 222- 230.

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