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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 summa­ rized. 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, Lan­ kester Botanical Garden, Universidad de Costa Rica INTRODUCTION The cohesion of pollen into compact units (the pollinia) and the evolution, in the Orchida­ ceae, of accessory structures for attachment to pollinators, has attracted the attention of scien­ tists since the time of Darwin, whofound orchid poIlinaria to be "as perfect as any of the most beautiful adaptations in the animal kingdom" (Darwin 1877). If the grouping of pollen grains into tetrads, reduction in exine surrounding inner grains, and pollen consolidation into pollinia are also known in the Asclepiadaceae, only the Or­ chidaceae show the innovation of pollinaria . This term refers to the entire package, including one or more pollinia and the associate structures, which is removed by pollinators, and its evolu­ tion is a key to explain sorne of the major mor­ phological differences between orchids' ftowers and those of other plant farnilies. * Corresponding author. In a family whose ftowers are pollinated mostly by insects, powdery pollen (which is more suited for the placement on hairy parts) was less effective in promoting dispersal, and the evolution of an adhesive viscidium was cru­ cial to maxirnize the efficiency of pollen export through the improvement of its adherence to smooth surfaces such as an insect's thorax, mouthparts, frons, eyes, and legs (Dressler 1981). The efficiency of pollen transfer depends not only on the selection of visitors by means of specific visual and olfactory cues, but also on the precise deposition of the pollen masses on a specific target area of the pollinator's body. This • involves modifications in perianth parts as weIl as in gynoecial and androecial organs, and even­ tually the shape and size of the viscidium. The precision of pollen placement on specific parts of the body of pollinators ensures interspecific pollination and, on the other side, allows several sympatric orchids to share the same pollinators without interference in pollen deposition (Dress- 1er 1968, Nilsson 1983, Steiner 1989). Mechan- 69 70 SELBYANA Volume 29(1) 2008 FiGURE 1. Pollinaria in dorsal and ventral views. A. Acineta densa (JBL-09182). B. unidentified Acineta species (Pupulin 6291). Scale bar = 5 mm. ical movements of the caudicles and stipe after pollinarium removal re-orient the pollinia and maximize pollen distribution. Bending move­ ments 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 stig­ matic entrance (Borba & Semir 1999), prevent geitonogamy and promote effective outcrossing in pollen transfer. The high degree of variation in pollinaria or­ ganization and morphology reflects the central role of this structure in orchid pollination mech­ anisms, and Johnson and Edwards (2000) sug­ gested it was pivotaI in promoting the tremen­ dous radiation of the Orchidaceae, which have more than 25,000 species (Atwood 1986, Cribb & Govaerts 2005). The characteristics of pollinia and the asso­ ciated structures have been used as a tool for orchid classification since the early work of John Lindley; Reichenbach (1852) devoted his doc- FIGURE 2. PoUinaria in dorsal, ventral, and late ral views. A. Kefersteinia parvilabris (Bogarfn 806). B. Chondroscaphe atrilinguis (Bogarfn 2950). Scale bars = 1 mm. toral dissertation to the study of poIlinaria as in­ dicators 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 massu­ lae), as well as the structure, development, or­ ganization and shape of poIlinaria (including the presence of caudicles and their composition, and the presence, number and shape of stipes), have been employed as sources of characters at vari­ ous systematic levels to define taxa from species to subfamilies. The use of pollinaria information as an addition al tool in taxonomic work is a cur­ rent practice at Lankester Botanical Garden (LBG), where researchers make use of their morphological features in a broad range of sys­ tematic 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 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 71 FIGURE 3. A. Cases hosting the collection of pol­ linaria. B. Envelopes to store pollinaria in the collec­ tion of Lankester Botanical Garden. construction of phylogenetic hypotheses (i.e., Williams 1970a, 1970b, 1972; Ackerman & Wil­ liams 1981, Burns- Balogh 1982, Dressler 1986, 1993; Chase 1987, Freudenstein & Rasmussen 1997). Accumulating evidence shows that floral evo­ lution in the Orchidaceae is essentially driven by pollinator traits , linking the study of pollination to evolutionary biology. New, intricate orchid­ pollinator interactions are continuously revealed through the application of accurate empirical studies that improve our understanding of plant evolution in general. However, direct observa­ tion of pollination acts is notoriously difficult, and the lack of direct records inhibits a better understanding of the pollination biology of man y orchid taxa, and particularly of the real degree of pollinator specificity. The study of mating patterns and male fitness in orchid repro­ duction is dependent on a precise identification of pollinaria. Pollinaria found on insects are of­ ten informative of the presence of rare species or of orchids that occupy difficult to reach hab­ itats 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 interest­ ing case of the study of orchid pollination with­ out any orchids, through the identification of or­ chid genera and species from the pollinaria borne by captured insects, stressing the impor­ tance of a reference collection to compare pol­ linaria. Recent work by Widmer and collabora­ tors addressed the problem of the relative pau­ city of field observations by means of a molec­ ular approach, based on the analysis of DNA recovered from pollinaria found on insects (Wid­ mer 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 pol­ linaria originated, disclosing previously unre­ corded orchid-pollinator relationships. ORCHID POLLINARIA COLLECTION AT LANKESTER BOTANICAL GARDEN To support studies in systematic botany and pollination biology, LBG began in 2003 an ef­ fort to build and maintain a large collection of orchid pollinaria, with emphasis on Neotropical taxa and special groups of interest to the re­ search staff. Pollinaria are mostly removed from plants grown in the living collection of the center, in­ c1uding more than 15 ,000 accessions belonging to approximately 1300 species. Priority was giv­ en to Neotropical and Mesoamerican genera and species, and to the groups that exhibit potentially informative pollinarium morphology, with em­ phasis on taxa with highly specialized reproduc­ tive biology-" It is not insignificant that the most­ ly 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 polli­ naria intended for the collection are removed with a thin-pointed instrument, trying to mini- 72 SELBYANA Volume 29(1) 2008 TABLE 1. Orchid pollinaria in the collection of Lankester Botanical Garden, University of Costa Rica. No. Genus/species Country Collector Date P0340 Acineta Costa Rica 14.11.2006 P0359 Acineta Costa Rica 14.11.2006 P0347 Acineta Costa Rica Pupulin 6291 14.11.2006 P0362 A cineta densa Costa Rica 14.11.2006 P0303 Acineta densa Costa Rica G6mez 7 13.09.2006 P0358 Acineta densa Costa Rica Pupulin 4200 14.11.2006 POl17 Acineta densa . Costa Rica 09.06.2005 P0118 Acineta densa Costa Rica 09.06.2005 P0119 Acineta densa Costa Rica 09.06.2005 P0120 Acineta densa Costa Rica Pupulin 4200 09.06.2005 P0467 Ada aurantiaca Colombia 20.07.2007 P0403 Ada aurantiaca Colombia 19.07.2007 P0086 Ada chlorops Costa Rica 13.05.2005 P0278 Aerides falcata Thailand 01.09.2006 P0171 Aerides multifiora Thailand 07 .07.2005 POO13 Angraecum eburneum Madagascar 15.12.2003 P0121 Angraecum eichlerianum Africa 09.06.2005 P0336 Arpophyllum giganteum Costa Rica 26.10.2006 P0071 Ascocentrum curvifolium Thailand 13.05.2005 P0087 Aspasia epidendroides Costa Rica Vargas s.n. 13.05.2005 P0122 Barkeria lindleyana Costa Rica 09.06.2005 P0246 Batemania leferenzii Peru Pupulin 6224 13.06.2006 P0405 Bensteinia ramonensis Costa Rica Bogarin 1923 19.07.2007 P0194 Bensteinia ramonensis Ecuador Bogarin 1923 17.03.2006 P0224 Benzingia cornuta Ecuador Pupulin 5989 16.05.2006 P0411 Benzingia reichenbachiana Costa Rica Bogarin J 890 19.07.2007 P0153 Benzingia caudata Ecuador Pupulin 5989 19.04.2006 P0045 Benzingia reichenbachiana Costa Rica Whitten 2052 13.05.2005 P0123 Brassavola nodosa Costa Rica 09.06.2005 P0450 Brassia arcuigera Costa Rica Bogarin 2416 20.07.2007 P0456 Brassia arcuigera Costa Rica Bogarin 1640 20.07.2007 P0149 Brassia arcuigera Costa Rica Pupulin 4886 14.06.2005 POl86 Brassia arcuigera Costa Rica Pupulin 5646 22.12.2005 P0404 Brassia arcuigera Costa Rica Bogarin 2416 19.07.2007 P0468 Brassia caudata Costa Rica Bogarin 2554 20.07.2007 P0021 Brassia caudata Costa Rica 22.10.2005 P0088 Brassia gireoudiana Costa Rica 13.05.2005 P0188 Brassia gireoudiana Costa Rica 22.1 2.2005 P0261 Brassia suavissima Costa Rica Bogarin 2855 30.06.2006 P0150 Brassia verrucosa Costa Rica 14.06.2005 P0170 Brassia caudata Costa Rica Herra J 07.07.2005 POI64 Brassia caudata Jamaica Hamlyn s.n. 07.07 .2005 POO05 Brassia arcuigera Costa Rica Pupulin s. n. 06.08.2003 P0386 Bulbophyllum lobbii Thailand 07.03.2006 P0302 Catasetum maculatum Costa Rica Bogarin 2609 13.09.2006 POOOI Catasetum maculatum Costa Rica Elizondo s.n. 20.07 .2004 P0354 Catasetum maculatum Costa Rica Rojas 137 14.11.2006 P0124 Cattleya intermedia Brazil 09.06.2005 P0381 Chaubardia heteroclita Peru Pupulin 6518 05.03.2006 P0287 Chaubardiella pubescens Costa Rica Pupulin 6240 05 .09 .2006 P0308 Chaubardiella subquadrata Costa Rica 20.09.2006 P0044 Chaubardiella pacuarensis Costa Rica 13.05.2005 P0042 Chaubardiella pacuarensis Costa Rica Blanco 1072 13.05.2005 P0282 Chondrorhyncha thienii Colombia Pupulin 6281 05.09.2006 P0481 Chondroryncha thienii Ecuador 19.08.2007 P0380 Chondroscaphe amabilis Ecuador Pupulin 6513 05.03.2006 P0376 Chondroscaphe atrilinguis Panama Bogarin 2950 12.03.2006 P0409 Chondroscaphe atrilinguis Panama Bogarin 2950 19.07.2007 P0465 Chondroscaphe bicolor Costa Rica Pupulin 3508 20.07.2007 P0396 Chondroscaphe bicolor Costa Rica 12.03.2006 P0339 Chondroscaphe bicolor Costa Rica Bogarin 1894 14.11.2006 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 73 TABLE l. Continued. No. Genus/species Country Collector Date P0341 Chondroscaphe bicolor Costa Rica Bogar[n 2203 14.11.2006 P0326 Chondroscaphe bicolor Panama 03.10.2006 P0408 Chondroscaphe bicolor Costa Rica Bogar[n 1914 19.07.2007 P0410 Chondroscaphe bicolor Costa Rica Bogar[n 996 19.07.2007 P0412 Chondroscaphe bicolor Costa Rica Bogar[n 426 19.07.2007 P0413 Chondroscaphe bicolor Costa Rica Bogarin 2874 19.07.2007 P0429 Chondroscaphe bicolor Costa Rica Bogarin 984 19.07.2007 P0355 Chondroscaphe bicolor Costa Rica Hoffman s.n. 14.11.2006 P0050 Chondroscaphe bicolor Costa Rica 13.05.2005 P0049 Chondroscaphe bicolor Costa Rica 13.05.2005 P0048 Chondroscaphe bicolor Costa Rica Pupulin 3525 13.05.2005 P0416 Chondroscaphe bicolor Costa Rica Pupulin 3525 19.07.2007 P0168 Chondroscaphe bicolor Costa Rica Bianchi 68 07.07.2005 P0346 Chondroscaphe bicolor Costa Rica Bianchi 68 14.11.2006 P0394 Chondroscaphe bicolor Costa Rica Bogarin 996 12.03.2006 P0342 Chondroscaphe bicolor Costa Rica Karremans 883 14.11.2006 P0473 Chondroscaphe eburnea Panama Dressler 6826 19.08.2007 P0439 Chondroscaphe eburnea Panama Dressler 6826 20.07.2007 P0379 Chondroscaphe embreei Ecuador Pupulin 6510 05.03.2006 P0464 Chondroscaphe embreei Ecuador Pupulin 6510 20.07.2007 P0329 Chondroscaphe escobariana Ecuador Pupulin 6289 17.10.2006 P0469 Chondroscaphe flaveola Ecuador Pupulin 6474 20.07.2007 P0378 Chondroscaphe flaveola Ecuador Pupulin 6474 05.03.2006 P0425 Chondroscaphe flaveola Ecuador Pupulin 6474 19.07.2007 P0325 Chondroscaphe yamilethiae Costa Rica Pupulin 4637 03.10.2006 P0398 Chondroscaphe yamilethiae Costa Rica Pupulin 4637 12.03.2006 P0322 Chondroscaphe yamilethiae Costa Rica Pupulin 4701 03.10.2006 POllO Chondroscaphe yamilethiae Costa Rica Pupulin 4638 24.05.2003 P0047 Chondroscaphe yamilethiae Costa Rica Pupulin 4637 13.05.2005 P0125 Coccineorchis standleyi Costa Rica Pupulin 4229 09.06.2005 P0156 Cochleanthes aromatica Costa Rica 16.05.2005 P0067 Cochleanthes aromatica Costa Rica 13.05.2005 P0126 Coelia bella Guatemala (?) 09.06.2005 P0185 Coeliopsis hyacinthosma Costa Rica Pupulin 4477 22.12.2005 P0475 Coeliopsis hyacinthosma Costa Rica Pupulin 4377 19.08.2007 P0089 Comparettia falcata Costa Rica Aguilar J 13.05.2005 P0310 Cycnoches amparoanum Costa Rica Rojas 114 20.09.2006 P0352 Cymbidium iridioides India Pupulin 6416 14.11.2006 P0026 Cymbidium iridoiides India 22.10.2005 P0216 Cyrtopodium macrobulbon Costa Rica Salazar s.n. 03.05.2006 P0301 Daiotyla albicans Costa Rica Pupulin 4372 07.09.2006 P0479 Daiotyla albicans Costa Rica Pupulin 4372 19.08.2007 P0300 Daiotyla crassa Costa Rica Pupulin 4257 07.09.2006 POO03 Daiotyla crassa Costa Rica Pupulin 4257 18.03.2004 P0478 Daiotyla crassa Costa Rica Pupulin 4257 19.08.2007 P0428 Daiotyla xanthina Colombia Pupulin 5891 19.07.2007 P0388 Dendrobium antennatum New Guinea 07.03.2006 P0286 Dichaea Costa Rica Pupulin 5049 05.09.2006 P0037 Dichaea acroblephara Costa Rica Bogar{n 1009 13.05.2005 P0029 Dichaea acroblephara Costa Rica Pupulin 4945 13.05.2005 P0030 Dichaea acroblephara Costa Rica Pupulin 5049 13.05.2005 P0069 Dichaea amparoana Costa Rica Bogar[n 6 79 13.05.2005 P0272 Dichaea cryptarrhena Costa Rica Pupulin 5662 29.08.2006 P0275 Dichaea elliptica Costa Rica ~upulin 5133 0l.09.2006 P0038 Dichaea elliptica Costa Rica Pupulin 5133 13.05.2005 P0274 Dichaea fragrantissima subsp. eburnea Costa Rica Blanco 513 0l.09.2006 P0031 Dichaea fragrantissima subsp. eburnea Costa Rica Blanco 513 13.05.2005 P0032 Dichaea fragrantissima subsp. eburnea Costa Rica Pupulin 4601 13.05.2005 P0281 Dichaea globosa Costa Rica Pupulin 4517 05.09.2006 P0033 Dichaea globosa Costa Rica Pupulin 4517 13.05.2005 P0163 Dichaea globosa Costa Rica Klark 286 07.07.2005 74 SELBYANA Volume 29(1) 2008 TABLE 1. Continued . No. Genus/species Country Co llector Date P0027 Dichaea morrisii Costa Rica Whitten 2171 04.12.2003 P0284 Dichaea obovatipetala Costa Rica Pupulin 5059 05.09 .2006 POO1 2 Dichaea obovatipetala Costa Rica Pupulin 5059 13.01.2004 P0036 Dichaea panamensis Costa Rica Bogar[n 1075 13.05.2005 P0035 Dichaea panamensis Costa Rica Karremans 621 13.05.2005 P0040 Dichaea poicillantha Costa Rica Bogar[n 187 13.05.2005 P0039 Dichaea poicillantha Costa Rica Pupulin 4662 13.05.2005 P0406 Dichaea poicillantha Nicaragua Karremans 1140 19.07.2007 P0271 Dichaea trulla Costa Rica Pupulin 5130 29.08.2006 POl 74 Dichaea similis Costa Rica Pupulin 5644 17.07.2005 P0041 Dichaea similis Costa Rica Pupulin 4089 13.05.2005 P0I59 Dichaea trulla Costa Rica Blanco 2812 07.07.2005 P022J Dressleria d ilecta Costa R ica Pupulin 5156 09.05.2006 P0390 Encyclia ambigua Mexico 07 .03.2006 P0222 Encyclia cordigera Costa Rica 12.05.2006 pOln Encyclia ceratistes Costa Rica Pupulin 5520 17.07 .2005 P0025 Epidendrum Costa Rica 22.12.2005 P0043 Epidendrum Costa Rica 22.12.2005 P0046 Epidendrum Costa Rica Pupulin 5570 22.12.2005 P01 32 Epidendrum centradenia Costa Rica 09.06.2005 POO08 EpidendrL/m odontochilwn Costa Rica Montealegre 114 22.12.2005 P0I 31 Epidendrum schumannianL/m Costa Rica PupL/lin 4864 09.06.2005 POl 30 Epidendrum wallisii Costa Rica PL/pL/lin 3832 09 .06.2005 POJ27 Epigeneium cymbidioides Phi lippines 09.06.2005 P0488 Eriopsis wercklei Costa Rica 14.01.2008 P0489 Eriopsis werckle i Costa Rica 14.01.2008 POO06 Erycina crista-galli Costa Rica PL/pL/lin s.n. 25 .02.2005 P02 18 Erythrodes tridax Costa Rica Bogar[n 993 03.05 .2006 P0480 Euryblema anatonwn Panama 19.08.2007 P0307 Euryblema andreae Colombia PL/pU lin 6282 13.09.2006 P0299 Euryblema andreae Colombia PL/pulin 6282 06.09.2006 POl 66 Eurychone rothschildiana Africa 07 .07.2005 POl54 Galeandra lac us tris Ecuador 14.06.2005 P041 9 Galeottia grandiflora Costa Rica 19.07 .2007 POliS Galeottia grandiflora Costa Rica Whitten 2138 08.05.2004 P0060 Galeottia grandiflora Costa Rica Whitten 2058 13.05.2005 POI 96 Gastrochilus calceola ris Thailand 17.03.2006 POl 95 GastrochilL/S f ormosanum Asia 17.03.2006 P0447 Gongora Costa Rica Blanco 2609 20.07.2007 P0219 Gongora Costa Rica 03.05.2006 P0228 Gongora Costa Rica 17.05.2006 P046 1 Gongora Costa Rica Blanco 2598 20.07 .2007 P03 J9 Gongora Costa Rica Gômez 60 29.09.2006 P0255 Gongora Costa Rica Roj as 111 2 1.06.2006 P0317 Gongora Costa Rica 29.09.2006 P0312 Gongora arm eniaca Costa Rica Pupulin 5214 29.09.2006 P0436 Gongora armeniaca Costa Rica Pupulin 5758 20.07.2007 P0442 Gongora claviodora Costa R ica Serrano 125 20.07.2007 P0227 Gongora galeata Ecuador 17.05.2006 P03 13 Gongora galeata Mexico 29.09.2006 POOlS Gongora leucochila Costa Rica 19.01.2004 P0226 Gongora truncata Venezuela 17.05.2006 P0316 Gongora unicolor Costa Rica 29.09.2006 P03 11 Gongora unicolor Costa Rica Pupulin 4954 29.09.2006 P0229 Govenia ciliilabia Costa Rica PL/pulin 621 7 06.06.2006 P0332 Grammatophyllum multiflo rum Philippines 26.10.2006 POO17 GrammatophyliL/m scriptum Phi lippines 12.05.2004 P0493 Guarianthe skinneri Costa Rica 05.02.2008 POO14 Haraella retrocalla Taiwan 30.10.2002 POJ05 Heterotaxis sessilis Costa Rica Clark 209 J9.05.2005 P0324 Houlletia tigrina Costa Rica Bogar[n 2480 03 .10.2006 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 75 T ABLE 1. Continued. No. Genus/species Country Collector Date P0051 HuntLeya burtii Costa Rica 13.05.2005 P0402 Huntleya burtii Costa Rica 19.07.2007 POO92 Hygroehilus parishii Philippines 13 .05.2005 P0200 Kefersteinia Costa Rica Bogarfn 806 09.05.2006 P0197 Kefersteinia andreettae Ecuador 02.05.2006 P0395 Kefersteinia aurieuLata Panama DressLer 6831 12.03.2006 P0417 Kefersteinia eostarieensis Costa Rica Bogarfn 761 19.07.2007 P0056 Kefersteinia eostarieensis Costa Rica Pupulin 5029 13.05.2005 P0052 Kefersteinia exeentriea Costa Rica BLanco 1053 13.05.2005 P0426 Kefersteinia Laminata Ecuador Pupulin 655/ 19.07.2007 P0385 Kefersteinia orbieuLaris Costa Rica HoLst 8773 07.03.2006 P0155 Keferste inia orbieuLaris Costa Rica Pupulin s.n. 13.05.2005 P0057 Kefersteinia orbieuLaris Costa Rica PupuLin s.n. 13.05.2005 P0055 Kefersteinia parviLabris Costa Rica Bogarfn 143 13.05.2005 POl98 Keferste inia pellita Ecuador Pupulin 5998 02.05.2006 P023 1 Kefersteinia retanae Costa Rica Cambronero s.n. 06.06.2006 P0427 Kefersteinia taurina Ecuador Pupulin 6550 19.07.2007 P0193 Kefersteinia taurina Ecuador Pupulin 5892 17.03.2006 POO04 Kefersteinia toLùnensis Colombia Pupulin 4205 30.10.2002 P0420 Kefersteinia toLimensis Colombia Pupulin 4205 19.07 .2007 P0054 Kefersteinia tolimensis Colombia Pupulin 4205 13.05.2005 P0053 Kefersteinia werekLei Costa Rica PupuLin 3039 13.05.2005 P044 1 Koellensteinia graminea Ecuador PupuLin 6517 20.07.2007 P0165 LeoehiLus tricuspidatus Costa Rica PupuLin 5648 07.07.2005 P0128 LeucohyLe subuLata Nicaragua PupuLin 4301 09.06.2005 P0090 Loekhartia amoena Costa Rica PupuLin 4515 13.05 .2005 P0091 Loekhartia pittieri Costa Rica BLanco 2653 13.05.2005 P0184 Lyeasle Costa Rica 22. 12.2005 POl 87 Lyeaste Costa Rica 29.12.2005 P0217 Lyeaste Costa Rica 03 .05.2006 P0393 Lycaste Costa Rica 07.03.2006 P0454 Lyeaste Costa Rica 20.07.2007 P0462 Lyeaste Costa Rica Pupulin 2780 20.07.2007 P0444 Lyeaste deppei México- 20.07 .2007 Nicaragua P0230 Lycaste deppei México- 06.06.2006 Nicaragua P0460 Lyeaste Leueantha Costa Rica 20.07.2007 P0437 Lycaste leucantha Costa Rica Bogarfn 2070 20.07 .2007 P0024 Lyeaste Leucantha Costa Rica 03.10.2002 P0334 Lycaste x Michelii 26.10.2006 P0476 Lyeaste sehilleriana Panama 19.08.2007 P0201 Lycaste tricoLor Costa Rica 09.05 .2006 P0383 Lyeaste tricoLor Costa Rica 07.03.2006 P0391 Masdevallia reiehenbachiana Costa Rica Vega s.n. 07.03 .2006 P0129 Maxillaria Costa Rica 22.1 2.2005 P0180 Max illaria Costa Rica 22.12.2005 P0357 Maxillaria Costa Rica 14.11.2006 P0361 Maxillaria Costa Rica 14.11.2006 P0364 Maxillaria Costa Rica BLanco 2808 14.1 1.2006 P0235 Maxillaria Costa Rica BLanco 2808 06.06.2006 P0178 Maxillaria Costa Rica G6mez 40 22.1 2.2005 P0240 Maxillaria Costa Rica Pupulin 3023 13.06.2006 P0028 Maxillaria Costa Rica Whitten 2045 05 .05.2006 PO 107 Max illaria aeostae Costa Rica BLanco 941 19.05.2005 PO 103 Maxillaria aLba Costa Rica PupuLin 2850 19.05.2005 POIOI Maxillaria angustissima Costa Rica 19.05.2005 P0167 Maxillaria araehnitiflora Costa Rica Bogarfn 344 22.12.2005 P0262 Maxillaria araehnitifiora Costa Rica BLanco 2262 30.06.2006 P0095 Maxillaria atwoodiana Costa Rica 18.05.2005 P0234 Maxillaria atwoodiana Costa Rica 06.06.2006 POI04 Maxillaria braehybulbon Costa Rica Bogarfn 874 19.05.2005 76 SELBYANA Volume 29(1) 2008 TABLE 1. Continued. No. Genus/species Country Collector Date P0249 Maxillaria brunnea Costa Rica Whitten 2028 16.06.2006 P0189 Maxillaria caespitifica Costa Rica Blanco 1076 22.12.2005 P0268 Maxillaria campanulata Costa Rica Bogarfn 1801 12.07.2006 PO 179 Maxillaria campanulata Costa Rica Comez 48 22.12.2005 P0259 Maxillaria confusa Costa Rica Bogarfn 1324 30.06.2006 P0236 Maxillaria con/usa Costa Rica Whitten 2030 06.06.2006 P0365 Maxillaria con/usa Costa Rica Whitten 2030 14.1 1.2006 P0181 Maxillaria endresii Costa Rica Whitten 2089 22.12.2005 P0097 Maxillaria endresii Costa Rica 18.05.2005 P0264 Maxillaria haberi Costa Rica Bogar{n 1868 05.07.2006 P0093 Maxillaria hedwigae Costa Rica Pupulin J 859 18.05.2005 P0176 Maxillaria inaudita Costa Rica Comez S.n. 22.12.2005 P0106 Maxillaria lueri Costa Rica 19.05.2005 POO09 Maxillaria luteoalba Colombia 06.11.2002 P0244 Maxillaria meridensis Costa Rica Comez 19 13.06.2006 P0099 Maxillaria moralesii Costa Rica 18.05.2005 P0248 Maxillaria moralesii Costa Rica Pupulin 4682 16.06.2006 PO 102 Maxillaria nasuta Costa Rica Pupulin 3247 19.05.2005 PO 108 Maxillaria pseudoneglecta Costa Rica 19.05.2005 POI77 Maxillaria punctostriata Costa Rica Comez 1 22.12.2005 P0094 Maxillaria ramonensis Costa Rica 18.05.2005 P0242 Max!llaria reichenheimiana Costa Rica Blanco 2107 13.06.2006 POO96 Maxillaria reichenheimiana Costa Rica 18.05.2005 POOlO Maxillaria ringens Costa Rica 21.1 1.2002 P0366 Maxillaria ringens Costa Rica Comez 24 14.11.2006 POlOO Maxillaria ringens Costa Rica Comez 24 18.05.2005 P0098 Maxillaria rodrigueziana Costa Rica 18.05.2005 P0232 Maxillaria rodrigueziana Costa Rica 06.06.2006 P0239 Maxillaria sanguinea Costa Rica 07.06.2006 PO 109 Maxillaria schlechteriana Costa Rica Whitten 2/81 19.05.2005 POl57 Maxillaria scorpioidea Costa Rica 22.12.2005 P0321 Maxillaria serrulata Costa Rica Bogarfn 1135 03.10.2006 P024 1 Maxillaria tonduzii Costa Rica Blanco 3179 13.06.2006 P0243 Maxillaria wercklei Costa Rica Blanco 3179 13.06.2006 P0371 Mesospinidium horichii Costa Rica Pupulin 5357 25.01.2006 P0369 Mesospinidium panamense Panama Dressler s.n. 25.01.2006 P0370 Mesospinidium warszewiczii Costa Rica 25.01.2006 P0470 Micochilus bimentatus Costa Rica Pupulin 6690 13.08.2007 P0074 Miltonia regnellii Brazil 13.05.2005 P0080 Miltoniopsis phalaenopsis Colombia 13.05.2005 P0078 Miltoniopsis roezlii Colombia 13.05.2005 P0079 Miltoniopsis vexillaria Colombia 13.05.2005 POO77 Miltoniopsis warscewiczii Costa Rica 13.05.2005 P0382 Mormodes colossa Costa Rica Blanco 2691 07.03.2006 P0443 Mormodes fractiflexa Costa Rica Bogarfn 2705 20.07.2007 P0223 Mormodes horichii Costa Rica. Bogarfn 2561 12.05.2006 P0023 Mormodes atropurpurea Costa Rica 04.03.2005 P0190 Mormodes colossa Costa Rica Blanco 2192 22.12.2005 P0081 Mormolyca ringens Costa Rica 13.05.2005 P0215 Oncidium Costa. Rica 03.05.2006 P0214 Oncidium Costa Rica Comez 18 03.05.2006 P0034 Oncidium coquianum Costa Rica Pupulin 3438 19.04.2006 P0375 Oncidium dichromaticum Costa Rica 28.02.2006 P0434 Oncidium dichromaticum Costa. Rica Pupulin 2495 19.07.2007 P0373 Oncidium ensatum Ecua.dor Pupulin 6421 05.02.2006 P0372 Oncidium fuscatum Nicaragua Pupulin 6420 05.02.2006 P0374 Oncidium hyphaematicum Nicaragua Pupulin 6422 05.02.2006 P0432 Oncidium hyphaematicum N icaragua. Bogarfn 3793 19.07.2007 P0431 Oncidium kotzchianum Costa Rica Pupulin 5455 19.07.2007 P0331 Oncidium luteum Costa. Rica. Pupulin 6197 26.10.2006 P0182 Oncidium luteum Costa Rica Whitten 2192 22.12.2005 P01I3 Oncidium oliganthum Guatemala Pupulin 3378 24.05.2003 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 77 T AB LE 1. Continued. No. Genus/species Country Collector Date POIIl Oncidium parviflorum Costa Rica 24.05 .2003 P0458 Oncidium parviflorum Costa Rica Bogarfn 2468 20.07 .2007 POl83 Oncidium schroederianum Costa Rica 22.1 2.2005 P0440 Oncidium schroederianum Costa Rica Pupulin 6254 20.07.2007 P0418 Oncidium sp.nov. Costa Rica Karremans 1650 19.07.2007 P0333 Oncidium stenobulbon Costa Rica Bogar(n 1065 26.10.2006 POI14 Oncidium storkii Costa Rica 24.05.2003 P0250 Osmoglossum egertonii Costa Rica 16.06.2006 P0266 Otoglossum chiriquense Costa Rica Pupulin 6235 12.07.2006 P0082 Otoglossum globuliferum Costa Rica Pupulin 5296 13.05.2005 P0273 Paphinia subclausa Costa Rica Dressler 6590 01.09.2006 P0455 Paphinia subclausa Costa Rica Dressler 6590 20.07.2007 POl61 Paphinia subclausa Costa Rica Pupulin 4665 07.07.2005 P0446 Pescatorea cerina Costa Rfca Blanco 2642 20.07.2007 P0389 Pescatorea cerina Costa Rica Pupulin 2993 07.03.2006 P0068 Pescatorea cerina Costa Rica Pupulin 2097 17.07 .2005 P0061 Pescatorea cerina Costa Rica 13.05.2005 POO07 Pescatorea cerina Costa Rica 06.08.2003 P0062 Pescatorea dayana Colombia 13.05.2005 POI92 Phalaenopsis amabilis Asia Pupulin 5713 17.03 .2006 POO02 Phalaenopsis schilleriana Philippines 12.05 .2004 P0173 Phalaenopsis amabilis lndonesia Pupulin 5713 17.07.2005 P0397 Phalaenopsis luedemanniana Philippines 28.03.2006 POO18 Phalaenopsis violacea Malasya 22.10.2005 P0472 Polycicnis barbata Costa Rica Bogarfn 1898 19.08.2007 P0022 Polycycnis barbata Costa Rica 04.03.2005 P0265 Polycycnis barbata Costa Rica Bogarfn 1898 12.07.2006 P0314 Polycycnis barbata Costa Rica Bogarfn 1845 29.09.2006 P0063 Prosthechea brassavolae Costa Rica 22.12.2005 P0392 Prosthechea chacaoensis Costa Rica 07.03 .2006 POl34 Prosthechea neglecta Costa Rica 10.06.2005 POl62 Prosthechea chacaoensis Costa Rica 07 .07.2005 P0133 Prosthechea vitellina Mexico 10.06.2005 P0116 Rodriguezia venusta Ecuador Pupulin 5281 08.06.2005 P0158 Rodriguezia venusta Ecuador 16.05 .2005 P0270 Rossioglossum schlieperianum Costa Rica Bogarfn 2851 18.07.2006 P0083 Rossioglossum schlieperianum Costa Rica 13 .05.2005 POll2 Rudolfiella aurantiaca Ecuador 24.05 .2003 P0135 Schomburgkia lueddemannii Costa Rica 13.06.2005 POOll Sedirea japonica Japan 12 .05 .2004 P0463 Sievekingia fimbriata Costa Rica Bogarfn 2813 20.07.2007 P0445 Sievekingia fimbriata Costa Rica 20.07.2007 P0136 Sieveking ia fimbriata Costa Rica 13.06.2005 P0209 Sievekingia fimbriata Costa Rica 03.05.2006 P0433 Sievekingia suavis Costa Rica Bogarfn 2575 19.07.2007 P0438 Sigmatostalix cardioglossa Costa Rica Pupulin 3499 20.07.2007 P0233 Sigmatostallx dulcineae Costa Rica Bogar(n 2079. 06.06.2006 P0356 Sobralia Costa Rica Aguilar 13 14.11.2006 P0338 Sobralia Costa Rica Bogarfn 913 14 .11.2006 P0306 Sobralia Costa Rica Dressler 6723 13.09.2006 P0337 Sobralia Costa, Rica Dressler 6621 14.11.2006 P0363 Sobralia Costa Rica Karremans 661 14.11.2006 P0260 Sobralia amabilis Costa Rica Dressler 6713 30.06.2006 P0367 Sobralia andreae Colombia 14.11.2006 P025 1 Sobralia hel/eri Costa Rica Dressler 6539 21.06.2006 P0492 Sobralia luteo la Costa Rica PupiAlin 2629 05 .02.2008 P0343 Sobralia macra Costa, Rica 14.11.2006 P0259 Sobralia macrophyl/a Costa Rica Casteifranco 155 30.06.2006 P0256 Sobralia quinata Costa, Rica Pupulin 3664 30.06.2006 P0160 Sobralia leuçoxantha Costa Rica Pupulin 4184 07.07.2005 P0448 Stanhopea Costa Rica 20.07 .2007 P0263 Stanhopea Costa Rica 05.07.2006 78 SELBYANA Volume 29(1) 2008 TABLE,1. Continued. No. Genus/species Country Collector Date P0267 Stanhopea Costa Rica 12.07.2006 P0294 Stanhopea Costa Rica 05.09.2006 P0430 Stanhopea Costa Rica 19.07.2007 P0452 Stanhopea Costa Rica 20.07.2007 P0277 Stanhopea Costa Rica Bogarfn 1639 01.09.2006 P0387 Stmihopea Costa Rica Bogarfn 757 07 .03.2006 P0276 Stanhopea Costa Rica Pupulin 2960 01.09.2006 P0291 Stanhopea Costa Rica Pupulin 2960 05 .09.2006 P0280 Stanhopea Costa Rica Rojas 112 05.09.2006 P0285 Stanhopea Ecuador Pupulin 6279 05.09.2006 P0292 Stanhopea Nicaragua Karremans 557 05.09.2006 P0210 Stanhopea Venezuela 03.05 .2006 P0257 Stanhopea Venezuela 30.06.2006 P0247 Stanhopea cfcostaricensis Costa Rica 13.06.2006 P0290 Stanhopea cirrhata Costa Rica Bogarfn 1021 05.09.2006 P0204 Stanhopea cirrhata Costa Rica Bogarfn 1021 03.05.2006 P0203 Stanhopea cirrhata Costa Rica Castelfranco 153 03.05.2006 P0295 Stanhopea cirrhata Costa Rica Pupulin 2843 05.09.2006 P0315 Stanhopea cirrhata Costa Rica Pup ulin 5827 29 .09.2006 P0485 Stanhopea confusa Costa Rica Flores S.n. 02.10.2007 P0466 Stanhopea costaricensis Costa Rica 20.07.2007 P0225 Stanhopea costaricensis Costa Rica 16.05.2006 P045 1 Stanhopea costaricensis Costa Rica 20.07.2007 P0457 Stanhopea costaricensis Costa Rica 20.07.2007 P0474 Stanhopea costaricensis Costa Rica 19.08.2007 P0205 Stanhopea costaricensis Costa R ica 03.05 .2006 P0368 Stanhopea costaricensis x cirrhata Costa Rica Rojas 113 26.11.2006 P0254 Stanhopea ecornuta Costa Rica Rojas 110 2 1.06.2006 P0283 Stanhopea ecornuta Costa Rica 05 .09.2006 P0288 Stanhopea ecornuta Costa R ica 05.09.2006 P0289 Stanhopea ecornuta Costa Rica 05.09.2006 P0137 Stanhopea ecornuta Costa Rica 13.06.2005 P0305 Stanhopea ecornuta Costa Rica 13.09.2006 P0471 Stanhopea ecornuta Costa Rica 19.08.2007 P0202 Stanhopea x fowlieana Costa Rica 03 .05.2006 P0253 Stanhopea x fowlieana Costa Rica 21.06.2006 P0252 Stanhopea gibbosa Costa Rica 21.06.2006 P0459 Stanhopea gibbosa Costa Rica 20.07.2007 P01 40 Stanhopea gibbosa Costa Rica 13.06.2005 P0449 Stanhopea grandiflora 20.07.2007 P0330 Stanhopea oculata Mexico Rojas 134 26.10.2006 P0345 Stanhopea oculata Mexico Rojas 135 14.11.2006 P0353 Stanhopea pulla Costa Rica Roj as 136 14. 11.2006 P0206 Stanhopea pulla Costa Rica 03.05.2006 P0208 Stanhopea pulla Costa Rica 03.05 .2006 P0020 Stanhopea pulla Costa Rica 21.06.2003 P0344 Stanhopea wardii Costa Rica Bogarfn 630 14.11.2006 P0328 Stanhopea wardii Costa Rica Dressler 6564 17.10.2006 P0335 Stanhopea wardii Costa Rica G6mez 51 26.10.2006 P0296 Stanhopea wardii Costa Rica Pupulin 4930 05 .09.2006 P0484 Stanhopea wardii Costa Rica Pupulin 4390 02.10.2007 POO16 Stanhopea wardii Costa Rica Pupulin 11 79 20.07.2004 P0146 Stanhopea wardii Costa Rica Pupulin 4930 13.06.2005 P0147 Stanhopea wardii Costa Rica Pupulin 1179 13.06.2005 P0207 Stanhopea wardii Costa Rica 03.05.2006 P0482 Stanhopea wardii Costa Rica 28 .09.2007 P0483 Stanhopea wardii Costa Rica 28.09.2007 P0487 Stanhopea wardii Costa Rica 25.10.2007 POO19 Stanhopea wardii Costa Rica 05 .08.2003 P0138 Stanhopea wardii Costa Rica 13.06.2005 P0139 Stanhopea wardii Costa Rica 13.06.2005 P014 1 Stanhopea wardii Costa R ica 13.06.2005 P0142 Stanhopea wardii Costa Rica 13.06.2005 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 79 TABLE 1. Continued. No. Genus/species Country Collector Dale P0144 Stanhopea wardii Costa Rica 13.06.2005 P0145 Stanhopea wardii Costa Rica 13.06.2005 P0293 Stanhopea warscewicziana Costa Rica 05.09.2006 P0297 Stanhopea warscewicziana Costa Rica 06.09.2006 P0304 Stanhopea warscewicziana Costa Rica 13.09.2006 P0327 Stanhopea warscewicziana Costa Rica 10.10.2006 P0143 Stanhopea warscewicziana Costa Rica 13.06.2005 P0269 Stenia Ecuador Pupulin 6239 18.07.2006 P0348 Stenia calceolaris Ecuador Pupulin 6332 14.1 1.2006 P0350 Stenia dodsoniana Ecuador Pupulin 6340 14.11.2006 P0279 Stenia guttata Ecuador Pupulin 5937 05.09.2006 P0318 Stenia pallida Ecuador Pupulin 6286 29.09.2006 P0415 Stenia pallida Ecuador 19.07 .2007 P0191 Stenia stenioides Ecuador Pupulin 5893 17.03.2006 P0407 Ste nia stenioides Ecuador Pupulin 5893 19.07.2007 P0199 Stenotyla lankesteriana Costa Rica 02.05 .2006 POl52 Stenotyla lankesteriana Costa Rica 19.04.2006 P0066 Stenotyla lendyana Guatemala Hubein s.n. 13.05.2005 P0065 Stenotyla lendyana Guatemala 13.05.2005 P0421 Stenotyla lendyana Guatemala 19.07.2007 P0245 Stenotyla panamensis Panama Pupulin 5868 13.06.2006 P0477 Stenotyla panamensis Panama Pupulin 5869 19.08.2007 P0064 Stenotyla picta Costa Rica Pupulin 3031 13.05.2005 P0422 Stenotyla picta Costa Rica 19.07.2007 P0423 Stenotyla picta Costa Rica 19.07.2007 P0384 Stenotyla picta Costa Rica Dressler s.n. 07.03.2006 P0435 Stenotyla picta Panama Bogarfn 2960 19.07.2007 P0298 TeLipogon biolleyi Costa Rica Karremans 1317 06.09.2006 P0237 Telipogon costaricensis Costa Rica Rojas 7148 06.06.2006 P0238 Telipogon monticola Costa Rica Rojas 7157 06.06.2006 P0148 TeLipogon ampliflorus Costa Rica 13.06.2005 P0073 Ticoglossum krameri Costa Rica 13.05.2005 P0070 Ticoglossum oerstedii Costa Rica 13.05.2005 P0220 Tolumnia triquetra Jamaica 03.05.2006 P0323 Trevoria zalhbrukneriana Costa Rica Bogarfn 1900 03 .10.2006 P0211 Trevoria zalhbrukneriana Costa Rica G6mez 39 03.05.2006 P0085 Trichocentrum capistratum Costa Rica Pupulin 2075 13.05.2005 P0084 Trichocentrum cymbiglossum Costa Rica PupuLin 4754 13.05 .2005 P0377 Trichocentrum pfavii Costa Rica Pupulin 5826 05.03.2006 P0360 Trichoglottis Philippines 14.11.2006 P0424 Trichopilia Colombia Dressler 6807 19.07.2007 P0349 TrichopiLia Colombia PupuLin 6333 14.11.2006 P0490 Trichopilia Costa Rica Pupulin 641 8 14.01.2008 P0399 Trichopilia eneidae Panama 28.03.2006 P0491 Trichopilia fragrans Colombia 14.01 .2008 P0486 Trichopilia fragrans Colombia Pupulin 6341 25 .10.2007 P0351 Trichopilia aff. fra grans Colombia Pupulin 6341 14.11 .2006 POl69 Trichopilia galeottiana Costa Rica Bogarfn 145 07.07 .2005 P0400 Trichopilia hennisiana Colombia 28.03 .2006 POl51 Trichopilia marg inata Costa Rica 14.06.2005 P0076 Trichopilia marg inata Costa Rica Pupulin 4382 13.05 .2005 P0175 Trichopilia marginata Costa Rica Pupulin 4433 17.07.2005 P0309 Trichopilia nobilis Colombia 20.09.2006 P0453 Trichopilia olmosii Panama Bogarfn 2948 20.07 .2007 P0075 Trichopilia punicea Costa Rica Pupulin 5213 13.05 .2005 poon Trichopilia punicea Costa Rica 13.05.2005 P0320 Trichopilia turrialbae Costa Rica 03. l0.2006 P021 2 Vanda coerulea Philippines 03 .05 .2006 P0213 Vanda lamellata Philippines 03.05.2006 P04 4 Warczewiczella discolor Costa Rica 19.07.2007 P0059 Warscewiczella discolor Costa Rica 13.05.2005 P0058 Warczewiczella ionoleuca Ecuador PupuLin 5235 13.05.2005 P0401 Warczewiczella ionoleuca Ecuador PupuLin 5235 19.07.2007 80 Record: 470 Total: 493 Unsorted Record: .., Total: 493 Un50rted SELBYANA Volume 29(1) 2008 RASTREO ( SCAt> IONUf,I,srn P0470 JBL-INVENT JBL-s .n . FAMI:..Y Orchidaceae '5PECIES bimentotus GENUS Mfcrochilus INflIASP COUtfTRY Costa Rica . COLlDATE 30 .07.2007. COLLECTOR F. Pupulln COLLNUH.8ER 6690. TAXONOfl Det . F. Pupulin, 8.2007. DATADATE 13.08.2007. POll.MAT JBL-PoIIColl. HERSM.\T FAMAT JBl-Splrlt. ILWSTR.'AAT 1II1i\l~r\idad d~ Co ... ta Rica RASTREO (S(AN IDNUI4ER P0492 GENUS Sabra/ia (QUtfTRY Costa Rica . jBL·I~NENT JBL- 09442 INfRASP FAMlLY Orchidaceae SPECIES /uteolo 10.12.2000. COlLECTOR: F. Pupulin COlLNIJ.'.\8E:R 2629 . TAXONOE'T Det. R. l. Dressler,03.2007. POLlMAT JBl-polleaU. HERaMAT 'MAAT DATADATE 05.2.2008. ILlUSTRMAT Foto. FIGURE 4. Two pages from the pollinaria database at Lankester Botanical Garden. PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 81 ....... ·ea.... ..... ·~· .......... _· ... *' ... _,-. ... _, ..... !_·. __ ; ___ .. _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 caudi­ cles. 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 Ep­ son Perfection V200 Photo) at 2400 to 3200 dpi resolution, and saved as TIFF files including rel­ ative scales. Dorsal and ventral views are re­ corded (FIGURE 1), and a lateral view is included when feasible (FIGURE 2). The pollinaria are then stored in specifically designed white paper en­ velopes and kept in hermetic plastic boxes (FIG­ URE 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 refer­ ences in a database designed for that purpose (FileMaker Pro 9.0) (FIGURE 4). This includes locality and collecting data, as weil as a cross­ reference 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 da­ tabase 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 (sub­ tribes Cranichidinae and Spiranthinae), the com­ pld pollinaria of the Epidendroideae are the best represented. In this subfarnily, we have rep­ resentative species of the Sobralieae (13 polli­ naria), Calypsoeae (1), Epidendreae (23), Cym­ bidieae (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 On­ cidiinae (89 pollinaria), Maxillarinae (69) , Stan­ hopeinae (110), and Zygopetalinae (149). Also present are pollinaria of Paleotropical Epiden­ droideae, 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 tenden­ cies in the organization of pollen and associate structures in this subtribe (Pupulin in prep.), and designed to use pollinaria morphology for spe­ cies identification in the genus Stanhopea (FIG­ URE 7). ACKNOWLEDGMENTS We are indebted to all the colleagues at LBG who helped through the years documenting pol­ linaria for collection, and in particular to Diego Bogarfn, Gustavo Rojas, Hilda Le6n-Paez, Jai­ me Aguilar, and Emily Serrano. We thank Margherita Pupulin and Carlotta Pupulin, who devoted part of their summer holidays to sub­ stantially improve the collection. All the polli­ naria included in the collection were removed from plants collected in the field or obtained through scientific interchange, thanks to scien­ tific 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 Collec­ tions 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); 2- D. 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. flaveola (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. reichenbachi­ ana (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). 36- 41 : Chaubardiella. 36-C pubescens (Pupulin 6240); 37-C aff. pubescens (Pupulin 6566); 38-C subquad­ rata (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 flabelliformis (Pupulin 6621). 66: Zyg­ osepalum lindeniae (Pupulin 6643). 67: Ixyophora viridisepala (Pupulin 6554). 68: "Chondrorhycnha " velas­ tiguii (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). 75- 78: 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 species. 1- 16: S. wardii. I- Dressler 6564; 2 - JBL-04360; 3 - JBL- 06666; 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. 17- 21: S. warscewicziana.17 - JBL-14610; 18 - JBL-08955; 19 - JBL-06148; 20 - JBL-14605 ; 21 - JBL-07747. 22- 28: 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 - JBL­ JJ696; 33 - JBL-06132 ; 34 - Rojas 110; 35 - JBL-11698. 36- 39: S. pulla. 36 - JBL-05657; 37 PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 85 TABLE 2. Polli naria included in the collection, ar­ ranged 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. Orchidoideae Cranichideae Goodyerinae: Erythrodes tridax, Microchilus bi­ mentatus Spiranthinae: Coccineorchis standleyi Epidendroideae Sobralieae: Sobralia amabilis, S. andreae, S. hel­ leri, S. luteola, S. mac ra, S. macrophylla, S. quinata, S. leucoxantha, Sobralia ssp. (5) Calypsoeae: Govenia ciliilabia Epidendreae Pleurothallidinae: Masdevallia reichenbachiana Laeliinae : Arpophyllum giganteum, Barkeria lin­ dleyana, Brassavola nodosa, Cattleya inlerme­ dia, Encyclia ambigua, E. cordigera, E. cera­ tistes, Epidendrum centradenia, E. odontochilum, E. schumannianum, E. wallisii, Epidendrum spp. (3), Guarianthe skinneri, Prosthechea brassavolae, P. chacaoensis (2), P. neglecta, P. vitellina, Schomburgkia luedde­ mannii Coeliinae [?): Coelia bella Cymbidieae f-- Catasetinae: Catasetum maculatum (3), Cycnoch­ es amparoanum, Cyrtopodium macrobulbon, Dressleria dilecta, Galeandra lacustris, Mor­ modes atropurpurea, M. colossa (2), M. fra cti­ flexa, M. horichii Cymbidiinae: Cymbidium iridioides (2), Gramma­ tophyllum multiflorum, G. scriptum Eriops idinae: Eriopsis wercklei (2) Oncidiinae: Ada aurantiaca (2), A. chlorops, Aspa­ sia epidendroides, Brassia arcuigera (6), B. cau­ data (4), B. gireoudiana (2), B. suavissima, B. verrucosa, Comparettia falcata, Erycina crista­ galli, Leochilus tricuspidatus, Lockhartia amoena, L. pittieri, Mesospinidium horichii, M. pana­ mense, M. warszewiczii, Miltonia regnellii, Milto­ niopsis phalaenopsis, M. roezlii, M. vexillaria, M. warscewiczii, Oncidium coquianum, O. dichroma­ ticum, O. dichromaticwn, 0. ensatum, O. fusca­ tum, O. hyphaematicum (2), 0. klotzchianum, O. luteum (2), O. oliganthum, O. parviflorum (2), O. parviflorum, O. schroederianum (2), 0. stenobul­ bon, O. storkii, Oncidium spp. (3), Osnwglossum egertonii, Otoglossum chiriquense, 0. globulifer­ um, Rodriguezia venusta (2), Rossioglossum schlieperianum (2), Sigmatostalix dulcineae, S. cardioglossa, Telipogon ampliflorum, T. biolleyi, T. costaricensis, T. monticola, Ticoglossum krameri, T. oerstedii, Tolumnia triquelra, Tricho­ cenlrum capistratum, T. cymbiglossum, T. pfavii, TABLE 2. Continued. Trichopilia eneidae, T. fragans (2), T. aff. fra­ grans, T. galeottiana, T. hennisiana, T. marginata (3), T. nobilis. T. olmosii, T. punicea (2), T. subu­ Lata, T. turrialbae, Trichopilia spp. (3) Maxillariinae: Helerotaxis sessilis, Lycasle borucana (2), L. deppei (2), L. leucantha (3), L. michelii (lassioglossa X cochleata), L. schilleriana, L. tri­ color (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. san­ guinea, M. schlechteriana, M . scorpioidea, M. ser­ rulata, 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 sub­ clausa (3), Polycicnis barbata (4), Sieveking iajim­ briata (3), S. sua vis, Stanhopea cirrhata (5), S. confusa, S. costaricensis (6), S. cf. costaricensis, S. costaricensis X cirrhata, S. ecornuta (7) , S. gibbo­ sa (3), S. grandiflora, S. oculata (2), S. pulia (4), S. wardii (19) , S. warscewicziana (5) , S. X fow­ lieana (2), Stanhopea spp. (14), Trevoria zalhbruk­ neriana (2) Coeliopsidinae: Coeliopsis hyacinlhosma (2) Zygopetalinae: Batemania leferenzii, X Bensteinia ramonensis (2) , Benzing ia caudata, B. cornuta, B. reichenbachiana (2), Chaubardia heteroclita, Chaubardiella pubescens, C. subquadrata, C. pac­ uarensis (2), Chondroryncha thienii (2) , Chondros­ caphe 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. panamen­ sis (2), D. poicillantha (3), D. similis, D. cf. simi­ lis, D. trulla (2), Euryblema anatonum, E. andreae (2), Galeottia grandiflora (3), Huntleya burtii (2) , Kefersteinia andreettae, K. auriculata, K. costari­ censis (2), K. excentrica, K. laminata, K. orbicu­ Laris (2) , K. pan1ilabris, K. pellita, K. retanae, K. taurina(2), K. tolimensis (3), K. wercklei, Kefer­ steinia spp. (2), Koellensteinia g raminea, Pescato­ rea cerina (5), P. dayana, Stenic calceolaris, S. dodsoniana, S. guttata, Stenia pallida (2), Stenia stenoides (2), Stenotyla lankesteriana (2), S. len- (Continued) - JBL-06155; 38 - Rojas 136; 39 - JBL-07727. 40-44: S. cirrhata. 40 - DB 1021; 41- DB 1021; 42 - FP 2843; 43 - FP 583; 44 - Castelfranco 153. 45- 47: S. gibbosa. 45 - JBL-04956; 46 - JBL-07529; 47 - JBL-07529. 48 - S. confusa (JBL-09484). 49 - S. grandiflora (JBL-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 SELBYANA Volume 29(1) 2008 TABLE 2. Continued. dyana (3), S. panamensis, S. picta (5), Warczewiczella discolor (2), W. ionoleuca (2) Vandeae Aeridinae: Aerides falcata, A . multiflora, Asco cen­ trum curvifolium, Gastrochilus calceolaris, G. formosanum, Haraella retrocalla, Hygrochilus parishii, Phalaenopsis amabi[js (2), P. luedde­ manniana, 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, Dendrobium 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: Max­ illariinae) 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 Bulbophyllum: a different mechanism preventing self-pollination in Orchidaceae. Plant Syst. Evo\. 217 : 197-204. Chase, M.W. 1987. Systematic implications of polli­ narium morphology in Oncidium Sw., Odonto­ glossum 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 His­ tory Publications, Kota Kinabalu, Sabah. Cribb, P. and R . Govaerts. 2005. Just how many or­ chids are there? Pp. 161- 172 in A. Raynal­ Roques, 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 Mur­ ray, London. Dressler, R.L. 1968. Pollination by euglossine 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 Ge­ sellschaft, Frankfurt. 1981. The Orchids: Natural History and Clas­ sification. Harvard University Press, Cambridge, Mass. ---. 1986. Features of pollinaria Wld orchid clas­ sification. Lindleyana 1: 125- 130. ---. 1993. Phylogeny and ClassificatiQn of the Or­ chid Farnily. Timber Press, Portland. --- and W. KJilcunas. 2006. A Sectional Classifi­ cation of Trichopilia Lindley (Orchidaceae), Sel­ byana 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 carry­ over, 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 Dac­ tylorhiza sambucina (Orchidaceae). Bot. Notiser 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 reassess­ ment of Costa Rican Zygopetalinae (Orchida­ ceae). A systematic revision of Dichaea in Costa Rica. Harvard Pap. Bot. 12(1): 15- 154. ---. 2007b. Those violet things: the elusive Di­ chaeas of Costa Rica, an overview of Dichaea phylogeny and classification. 14'" EOCCE Pro­ ceedings. 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 sys­ tema ricligendis" Ph.D. diss. , Leipzig. Steiner, KE. 1989. The pollination of Disperis (Or­ chidaceae) by oil-collecting bees in southern Af­ rica. 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 pollinar­ ia in the Oncidiinae. Amer. Orch. Soc. Bull. 39: 32-43. ---. 1970b. Sorne observations on pollinaria in the Oncidiinae- II. Amer. Orch. Soc. Bull. 39: 207- 220. ---. 1972. Additional studies on pollinaria in the Oncidiinae. Amer. Orch. Soc. Bull. 41: 222- 230. 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