the orchid pollinaria collection at lankester botanical ... - Epidendra

Selbyana 29(1): 69- 86. 2008.<br />

THE ORCHID POLLINARIA COLLECTION AT LANKESTER BOTANICAL<br />

GARDEN, UNIVERSITY OF COSTA RICA<br />

FRANCO PUPULIN*<br />

Lankester Botanical Garden, University of Costa Rica. P.O. Box 1031-7050 Cartago,<br />

Costa Rica,. CA<br />

Angel Andreetta Research Center on Andean Orchids, University Alfredo Pérez Guerrero,<br />

Extension Gualaceo, Ecuador<br />

Harvard University Herbaria, Cambridge, MA, USA<br />

The Marie Selby Botanical Gardens, Sarasota, FL, USA<br />

Email: fpupulin@cariari.ucr.ac.cr<br />

ADAM KARREMANS<br />

Lankester Botanical Garden, University of Costa Rica. P.O. Box 1031-7050 Cartago,<br />

Costa Rica, CA<br />

Angel Andreetta Research Center on Andean Orchids, University Alfredo Pérez Guerrero,<br />

Extension Gualaceo, Ecuador<br />

ABSTRACT. The relevance of <strong>pollinaria</strong> study in <strong>orchid</strong> system<strong>at</strong>ics and reproductive biology is summarized.<br />

The Orchid Pollinaria Collection and <strong>the</strong> associ<strong>at</strong>e d<strong>at</strong>abase of Lankester Botanical Garden, University<br />

of Costa Rica, are presented. The <strong>collection</strong> includes 496 <strong>pollinaria</strong>, bèlonging to 312 species in 94 genera,<br />

with particular emphasis on Neotropical taxa of <strong>the</strong> tribe Cymbidieae (Epidendroideae). The associ<strong>at</strong>ed<br />

d<strong>at</strong>abase includes digital images of <strong>the</strong> <strong>pollinaria</strong> and is progressively made available to <strong>the</strong> general public<br />

through EPIDENDRA, <strong>the</strong> online taxonomic and nomencl<strong>at</strong>ural d<strong>at</strong>abase of Lankester Botanical Garden.<br />

Examples are given of <strong>the</strong> use of <strong>the</strong> <strong>pollinaria</strong> <strong>collection</strong> by researchers of <strong>the</strong> Center in a broad range of<br />

system<strong>at</strong>ic applic<strong>at</strong>ions.<br />

Key words: Orchid <strong>pollinaria</strong>, system<strong>at</strong>ic botany, pollin<strong>at</strong>ion biology, <strong>orchid</strong> <strong>pollinaria</strong> <strong>collection</strong>, Lankester<br />

Botanical Garden, Universidad de Costa Rica<br />

INTRODUCTION<br />

The cohesion of pollen into compact units<br />

(<strong>the</strong> pollinia) and <strong>the</strong> evolution, in <strong>the</strong> Orchidaceae,<br />

of accessory structures for <strong>at</strong>tachment to<br />

pollin<strong>at</strong>ors, has <strong>at</strong>tracted <strong>the</strong> <strong>at</strong>tention of scientists<br />

since <strong>the</strong> time of Darwin, whofound <strong>orchid</strong><br />

poIlinaria to be "as perfect as any of <strong>the</strong> most<br />

beautiful adapt<strong>at</strong>ions in <strong>the</strong> animal kingdom"<br />

(Darwin 1877). If <strong>the</strong> grouping of pollen grains<br />

into tetrads, reduction in exine surrounding inner<br />

grains, and pollen consolid<strong>at</strong>ion into pollinia are<br />

also known in <strong>the</strong> Asclepiadaceae, only <strong>the</strong> Orchidaceae<br />

show <strong>the</strong> innov<strong>at</strong>ion of <strong>pollinaria</strong> .<br />

This term refers to <strong>the</strong> entire package, including<br />

one or more pollinia and <strong>the</strong> associ<strong>at</strong>e structures,<br />

which is removed by pollin<strong>at</strong>ors, and its evolution<br />

is a key to explain sorne of <strong>the</strong> major morphological<br />

differences between <strong>orchid</strong>s' ftowers<br />

and those of o<strong>the</strong>r plant farnilies.<br />

* Corresponding author.<br />

In a family whose ftowers are pollin<strong>at</strong>ed<br />

mostly by insects, powdery pollen (which is<br />

more suited for <strong>the</strong> placement on hairy parts)<br />

was less effective in promoting dispersal, and<br />

<strong>the</strong> evolution of an adhesive viscidium was crucial<br />

to maxirnize <strong>the</strong> efficiency of pollen export<br />

through <strong>the</strong> improvement of its adherence to<br />

smooth surfaces such as an insect's thorax,<br />

mouthparts, frons, eyes, and legs (Dressler<br />

1981). The efficiency of pollen transfer depends<br />

not only on <strong>the</strong> selection of visitors by means of<br />

specific visual and olfactory cues, but also on<br />

<strong>the</strong> precise deposition of <strong>the</strong> pollen masses on a<br />

specific target area of <strong>the</strong> pollin<strong>at</strong>or's body. This<br />

• involves modific<strong>at</strong>ions in perianth parts as weIl<br />

as in gynoecial and androecial organs, and eventually<br />

<strong>the</strong> shape and size of <strong>the</strong> viscidium. The<br />

precision of pollen placement on specific parts<br />

of <strong>the</strong> body of pollin<strong>at</strong>ors ensures interspecific<br />

pollin<strong>at</strong>ion and, on <strong>the</strong> o<strong>the</strong>r side, allows several<br />

symp<strong>at</strong>ric <strong>orchid</strong>s to share <strong>the</strong> same pollin<strong>at</strong>ors<br />

without interference in pollen deposition (Dress-<br />

1er 1968, Nilsson 1983, Steiner 1989). Mechan-<br />

69

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70 SELBYANA Volume 29(1) 2008<br />

FiGURE 1. Pollinari a in dorsal and ventral views.<br />

A. Acineta densa (JBL-09182). B. unidentified Acineta<br />

species (Pupulin 6291). Scale bar = 5 mm.<br />

ical movements of <strong>the</strong> caudicles and stipe after<br />

pollinarium removal re-orient <strong>the</strong> pollinia and<br />

maximize pollen distribution. Bending movements<br />

of <strong>pollinaria</strong>, which correctly orient <strong>the</strong><br />

pollinia to contact <strong>the</strong> stigma (Darwin 1877,<br />

Nilsson 1980, Johnson & Nilsson 1999), as weIl<br />

as <strong>the</strong> shrinking of <strong>the</strong> pollinia to fit <strong>the</strong> stigm<strong>at</strong>ic<br />

entrance (Borba & Semir 1999), prevent<br />

geitonogamy and promote effective outcrossing<br />

in pollen transfer.<br />

The high degree of vari<strong>at</strong>ion in <strong>pollinaria</strong> organiz<strong>at</strong>ion<br />

and morphology reflects <strong>the</strong> central<br />

role of this structure in <strong>orchid</strong> pollin<strong>at</strong>ion mechanisms,<br />

and Johnson and Edwards (2000) suggested<br />

it was pivotaI in promoting <strong>the</strong> tremendous<br />

radi<strong>at</strong>ion of <strong>the</strong> Orchidaceae, which have<br />

more than 25,000 species (Atwood 1986, Cribb<br />

& Govaerts 2005).<br />

The characteristics of pollinia and <strong>the</strong> associ<strong>at</strong>ed<br />

structures have been used as a tool for<br />

<strong>orchid</strong> classific<strong>at</strong>ion since <strong>the</strong> early work of John<br />

Lindley; Reichenbach (1852) devoted his doc-<br />

FIGURE 2. PoUinaria in dorsal, ventral, and l<strong>at</strong>eral<br />

views. A. Kefersteinia parvilabris (Bogarfn 806). B.<br />

Chondroscaphe <strong>at</strong>rilinguis (Bogarfn 2950). Scale bars<br />

= 1 mm.<br />

toral dissert<strong>at</strong>ion to <strong>the</strong> study of poIlinaria as indic<strong>at</strong>ors<br />

of species rel<strong>at</strong>ionships. Diversity in<br />

number, shape, orient<strong>at</strong>ion, and packaging of<br />

pollinia (including <strong>the</strong> degree of tetrad packing,<br />

<strong>the</strong> st<strong>at</strong>e of <strong>the</strong> pollini a, <strong>the</strong> form, arrangement,<br />

orient<strong>at</strong>ion, and possible dimorphism of massulae),<br />

as well as <strong>the</strong> structure, development, organiz<strong>at</strong>ion<br />

and shape of poIlinaria (including <strong>the</strong><br />

presence of caudicles and <strong>the</strong>ir composition, and<br />

<strong>the</strong> presence, number and shape of stipes), have<br />

been employed as sources of characters <strong>at</strong> various<br />

system<strong>at</strong>ic levels to define taxa from species<br />

to subfamilies. The use of <strong>pollinaria</strong> inform<strong>at</strong>ion<br />

as an addition al tool in taxonomic work is a current<br />

practice <strong>at</strong> Lankester Botanical Garden<br />

(LBG), where researchers make use of <strong>the</strong>ir<br />

morphological fe<strong>at</strong>ures in a broad range of system<strong>at</strong>ic<br />

applic<strong>at</strong>ions (i.e., Dressler & Klikunas<br />

2006, Pupulin 2007a, 2007b; Bogarfn 2007,<br />

Pupulin in press). As <strong>pollinaria</strong> are less subject<br />

to parallelism than o<strong>the</strong>r floral fe<strong>at</strong>ures, <strong>the</strong>ir<br />

morphology also played a significant role in <strong>the</strong>

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PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 71<br />

FIGURE 3. A. Cases hosting <strong>the</strong> <strong>collection</strong> of <strong>pollinaria</strong>.<br />

B. Envelopes to store <strong>pollinaria</strong> in <strong>the</strong> <strong>collection</strong><br />

of Lankester Botanical Garden.<br />

construction of phylogenetic hypo<strong>the</strong>ses (i.e.,<br />

Williams 1970a, 1970b, 1972; Ackerman & Williams<br />

1981, Burns- Balogh 1982, Dressler 1986,<br />

1993; Chase 1987, Freudenstein & Rasmussen<br />

1997).<br />

Accumul<strong>at</strong>ing evidence shows th<strong>at</strong> floral evolution<br />

in <strong>the</strong> Orchidaceae is essentially driven by<br />

pollin<strong>at</strong>or traits, linking <strong>the</strong> study of pollin<strong>at</strong>ion<br />

to evolutionary biology. New, intric<strong>at</strong>e <strong>orchid</strong>pollin<strong>at</strong>or<br />

interactions are continuously revealed<br />

through <strong>the</strong> applic<strong>at</strong>ion of accur<strong>at</strong>e empirical<br />

studies th<strong>at</strong> improve our understanding of plant<br />

evolution in general. However, direct observ<strong>at</strong>ion<br />

of pollin<strong>at</strong>ion acts is notoriously difficult,<br />

and <strong>the</strong> lack of direct records inhibits a better<br />

understanding of <strong>the</strong> pollin<strong>at</strong>ion biology of<br />

man y <strong>orchid</strong> taxa, and particularly of <strong>the</strong> real<br />

degree of pollin<strong>at</strong>or specificity. The study of<br />

m<strong>at</strong>ing p<strong>at</strong>terns and male fitness in <strong>orchid</strong> reproduction<br />

is dependent on a precise identific<strong>at</strong>ion<br />

of <strong>pollinaria</strong>. Pollinaria found on insects are often<br />

inform<strong>at</strong>ive of <strong>the</strong> presence of rare species<br />

or of <strong>orchid</strong>s th<strong>at</strong> occupy difficult to reach habit<strong>at</strong>s<br />

and ecological niches. During a preliminary<br />

census of <strong>the</strong> <strong>orchid</strong> flora of <strong>the</strong> large «400<br />

hectares) Alberto M . Brenes Biological Reserve<br />

in Costa Rica, staff of LBG managed to record<br />

<strong>the</strong> presence of Houlletia tigrina on <strong>the</strong> basis of<br />

a single pollinarium found on an insect th<strong>at</strong> had<br />

been captured <strong>at</strong> <strong>the</strong> Reserve during a course on<br />

entomology. Dressler (1976) makes an interesting<br />

case of <strong>the</strong> study of <strong>orchid</strong> pollin<strong>at</strong>ion without<br />

any <strong>orchid</strong>s, through <strong>the</strong> identific<strong>at</strong>ion of <strong>orchid</strong><br />

genera and species from <strong>the</strong> <strong>pollinaria</strong><br />

borne by captured insects, stressing <strong>the</strong> importance<br />

of a reference <strong>collection</strong> to compare <strong>pollinaria</strong>.<br />

Recent work by Widmer and collabor<strong>at</strong>ors<br />

addressed <strong>the</strong> problem of <strong>the</strong> rel<strong>at</strong>ive paucity<br />

of field observ<strong>at</strong>ions by means of a molecular<br />

approach, based on <strong>the</strong> analysis of DNA<br />

recovered from <strong>pollinaria</strong> found on insects (Widmer<br />

et al. 2000). The study demonstr<strong>at</strong>es th<strong>at</strong> <strong>the</strong><br />

analysis of <strong>the</strong> nuclear ribosomal ITS region of<br />

<strong>the</strong> recovered pollen allows <strong>the</strong> identific<strong>at</strong>ion of<br />

<strong>the</strong> species or species-group from which <strong>the</strong> <strong>pollinaria</strong><br />

origin<strong>at</strong>ed, disclosing previously unrecorded<br />

<strong>orchid</strong>-pollin<strong>at</strong>or rel<strong>at</strong>ionships.<br />

ORCHID POLLINARIA COLLECTION AT<br />

LANKESTER BOTANICAL GARDEN<br />

To support studies in system<strong>at</strong>ic botany and<br />

pollin<strong>at</strong>ion biology, LBG began in 2003 an effort<br />

to build and maintain a large <strong>collection</strong> of<br />

<strong>orchid</strong> <strong>pollinaria</strong>, with emphasis on Neotropical<br />

taxa and special groups of interest to <strong>the</strong> research<br />

staff.<br />

Pollinaria are mostly removed from plants<br />

grown in <strong>the</strong> living <strong>collection</strong> of <strong>the</strong> center, inc1uding<br />

more than 15,000 accessions belonging<br />

to approxim<strong>at</strong>ely 1300 species. Priority was given<br />

to Neotropical and Mesoamerican genera and<br />

species, and to <strong>the</strong> groups th<strong>at</strong> exhibit potentially<br />

inform<strong>at</strong>ive pollinarium morphology, with emphasis<br />

on taxa with highly specialized reproductive<br />

biology-" It is not insignificant th<strong>at</strong> <strong>the</strong> mostly<br />

Neotropical advanced Cymbidieae account<br />

for almost 88% of <strong>the</strong> entire <strong>collection</strong>. Unlike<br />

<strong>the</strong> generalized use of mounting <strong>pollinaria</strong> on<br />

small paper triangles, to which <strong>the</strong>y are <strong>at</strong>tached<br />

by <strong>the</strong> n<strong>at</strong>ural glue of <strong>the</strong> viscidium, <strong>the</strong> <strong>pollinaria</strong><br />

intended for <strong>the</strong> <strong>collection</strong> are removed<br />

with a thin-pointed instrument, trying to mini-

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72 SELBYANA Volume 29(1) 2008<br />

TABLE 1. Orchid <strong>pollinaria</strong> in <strong>the</strong> <strong>collection</strong> of Lankester Botanical Garden, University of Costa Rica.<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

P0340 Acineta Costa Rica 14.11.2006<br />

P0359 Acineta Costa Rica 14.11.2006<br />

P0347 Acineta Costa Rica Pupulin 6291 14.11.2006<br />

P0362 A cineta densa Costa Rica 14.11.2006<br />

P0303 Acineta densa Costa Rica G6mez 7 13.09.2006<br />

P0358 Acineta densa Costa Rica Pupulin 4200 14.11.2006<br />

POl17 Acineta densa . Costa Rica 09.06.2005<br />

P0118 Acineta densa Costa Rica 09.06.2005<br />

P0119 Acineta densa Costa Rica 09.06.2005<br />

P0120 Acineta densa Costa Rica Pupulin 4200 09.06.2005<br />

P0467 Ada aurantiaca Colombia 20.07.2007<br />

P0403 Ada aurantiaca Colombia 19.07.2007<br />

P0086 Ada chlorops Costa Rica 13.05.2005<br />

P0278 Aerides falc<strong>at</strong>a Thailand 01.09.2006<br />

P0171 Aerides multifiora Thailand 07.07.2005<br />

POO13 Angraecum eburneum Madagascar 15.12.2003<br />

P0121 Angraecum eichlerianum Africa 09.06.2005<br />

P0336 Arpophyllum giganteum Costa Rica 26.10.2006<br />

P0071 Ascocentrum curvifolium Thailand 13.05.2005<br />

P0087 Aspasia epidendroides Costa Rica Vargas s.n. 13.05.2005<br />

P0122 Barkeria lindleyana Costa Rica 09.06.2005<br />

P0246 B<strong>at</strong>emania leferenzii Peru Pupulin 6224 13.06.2006<br />

P0405 Bensteinia ramonensis Costa Rica Bogarin 1923 19.07.2007<br />

P0194 Bensteinia ramonensis Ecuador Bogarin 1923 17.03.2006<br />

P0224 Benzingia cornuta Ecuador Pupulin 5989 16.05.2006<br />

P0411 Benzingia reichenbachiana Costa Rica Bogarin J 890 19.07.2007<br />

P0153 Benzingia caud<strong>at</strong>a Ecuador Pupulin 5989 19.04.2006<br />

P0045 Benzingia reichenbachiana Costa Rica Whitten 2052 13.05.2005<br />

P0123 Brassavola nodosa Costa Rica 09.06.2005<br />

P0450 Brassia arcuigera Costa Rica Bogarin 2416 20.07.2007<br />

P0456 Brassia arcuigera Costa Rica Bogarin 1640 20.07.2007<br />

P0149 Brassia arcuigera Costa Rica Pupulin 4886 14.06.2005<br />

POl86 Brassia arcuigera Costa Rica Pupulin 5646 22.12.2005<br />

P0404 Brassia arcuigera Costa Rica Bogarin 2416 19.07.2007<br />

P0468 Brassia caud<strong>at</strong>a Costa Rica Bogarin 2554 20.07.2007<br />

P0021 Brassia caud<strong>at</strong>a Costa Rica 22.10.2005<br />

P0088 Brassia gireoudiana Costa Rica 13.05.2005<br />

P0188 Brassia gireoudiana Costa Rica 22.1 2.2005<br />

P0261 Brassia suavissima Costa Rica Bogarin 2855 30.06.2006<br />

P0150 Brassia verrucosa Costa Rica 14.06.2005<br />

P0170 Brassia caud<strong>at</strong>a Costa Rica Herra J 07.07.2005<br />

POI64 Brassia caud<strong>at</strong>a Jamaica Hamlyn s.n. 07.07.2005<br />

POO05 Brassia arcuigera Costa Rica Pupulin s. n. 06.08.2003<br />

P0386 Bulbophyllum lobbii Thailand 07.03.2006<br />

P0302 C<strong>at</strong>asetum macul<strong>at</strong>um Costa Rica Bogarin 2609 13.09.2006<br />

POOOI C<strong>at</strong>asetum macul<strong>at</strong>um Costa Rica Elizondo s.n. 20.07.2004<br />

P0354 C<strong>at</strong>asetum macul<strong>at</strong>um Costa Rica Rojas 137 14.11.2006<br />

P0124 C<strong>at</strong>tleya intermedia Brazil 09.06.2005<br />

P0381 Chaubardia heteroclita Peru Pupulin 6518 05.03.2006<br />

P0287 Chaubardiella pubescens Costa Rica Pupulin 6240 05.09.2006<br />

P0308 Chaubardiella subquadr<strong>at</strong>a Costa Rica 20.09.2006<br />

P0044 Chaubardiella pacuarensis Costa Rica 13.05.2005<br />

P0042 Chaubardiella pacuarensis Costa Rica Blanco 1072 13.05.2005<br />

P0282 Chondrorhyncha thienii Colombia Pupulin 6281 05.09.2006<br />

P0481 Chondroryncha thienii Ecuador 19.08.2007<br />

P0380 Chondroscaphe amabilis Ecuador Pupulin 6513 05.03.2006<br />

P0376 Chondroscaphe <strong>at</strong>rilinguis Panama Bogarin 2950 12.03.2006<br />

P0409 Chondroscaphe <strong>at</strong>rilinguis Panama Bogarin 2950 19.07.2007<br />

P0465 Chondroscaphe bicolor Costa Rica Pupulin 3508 20.07.2007<br />

P0396 Chondroscaphe bicolor Costa Rica 12.03.2006<br />

P0339 Chondroscaphe bicolor Costa Rica Bogarin 1894 14.11.2006

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74 SELBYANA Volume 29(1) 2008<br />

TABLE 1. Continued.<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

P0027 Dichaea morrisii Costa Rica Whitten 2171 04.12.2003<br />

P0284 Dichaea obov<strong>at</strong>ipetala Costa Rica Pupulin 5059 05.09.2006<br />

POO1 2 Dichaea obov<strong>at</strong>ipetala Costa Rica Pupulin 5059 13.01.2004<br />

P0036 Dichaea panamensis Costa Rica Bogar[n 1075 13.05.2005<br />

P0035 Dichaea panamensis Costa Rica Karremans 621 13.05.2005<br />

P0040 Dichaea poicillantha Costa Rica Bogar[n 187 13.05.2005<br />

P0039 Dichaea poicillantha Costa Rica Pupulin 4662 13.05.2005<br />

P0406 Dichaea poicillantha Nicaragua Karremans 1140 19.07.2007<br />

P0271 Dichaea trulla Costa Rica Pupulin 5130 29.08.2006<br />

POl 74 Dichaea similis Costa Rica Pupulin 5644 17.07.2005<br />

P0041 Dichaea similis Costa Rica Pupulin 4089 13.05.2005<br />

P0I59 Dichaea trulla Costa Rica Blanco 2812 07.07.2005<br />

P022J Dressleria dilecta Costa R ica Pupulin 5156 09.05.2006<br />

P0390 Encyclia ambigua Mexico 07.03.2006<br />

P0222 Encyclia cordigera Costa Rica 12.05.2006<br />

pOln Encyclia cer<strong>at</strong>istes Costa Rica Pupulin 5520 17.07.2005<br />

P0025 Epidendrum Costa Rica 22.12.2005<br />

P0043 Epidendrum Costa Rica 22.12.2005<br />

P0046 Epidendrum Costa Rica Pupulin 5570 22.12.2005<br />

P01 32 Epidendrum centradenia Costa Rica 09.06.2005<br />

POO08 EpidendrL/m odontochilwn Costa Rica Montealegre 114 22.12.2005<br />

P0I 31 Epidendrum schumannianL/m Costa Rica PupL/lin 4864 09.06.2005<br />

POl 30 Epidendrum wallisii Costa Rica PL/pL/lin 3832 09.06.2005<br />

POJ27 Epigeneium cymbidioides Philippines 09.06.2005<br />

P0488 Eriopsis wercklei Costa Rica 14.01.2008<br />

P0489 Eriopsis wercklei Costa Rica 14.01.2008<br />

POO06 Erycina crista-galli Costa Rica PL/pL/lin s.n. 25 .02.2005<br />

P02 18 Erythrodes tridax Costa Rica Bogar[n 993 03.05.2006<br />

P0480 Euryblema an<strong>at</strong>onwn Panama 19.08.2007<br />

P0307 Euryblema andreae Colombia PL/pU lin 6282 13.09.2006<br />

P0299 Euryblema andreae Colombia PL/pulin 6282 06.09.2006<br />

POl 66 Eurychone rothschildiana Africa 07.07.2005<br />

POl54 Galeandra lac us tris Ecuador 14.06.2005<br />

P041 9 Galeottia grandiflora Costa Rica 19.07.2007<br />

POliS Galeottia grandiflora Costa Rica Whitten 2138 08.05.2004<br />

P0060 Galeottia grandiflora Costa Rica Whitten 2058 13.05.2005<br />

POI 96 Gastrochilus calceola ris Thailand 17.03.2006<br />

POl 95 GastrochilL/S f ormosanum Asia 17.03.2006<br />

P0447 Gongora Costa Rica Blanco 2609 20.07.2007<br />

P0219 Gongora Costa Rica 03.05.2006<br />

P0228 Gongora Costa Rica 17.05.2006<br />

P046 1 Gongora Costa Rica Blanco 2598 20.07.2007<br />

P03 J9 Gongora Costa Rica Gômez 60 29.09.2006<br />

P0255 Gongora Costa Rica Rojas 111 2 1.06.2006<br />

P0317 Gongora Costa Rica 29.09.2006<br />

P0312 Gongora arm eniaca Costa Rica Pupulin 5214 29.09.2006<br />

P0436 Gongora armeniaca Costa Rica Pupulin 5758 20.07.2007<br />

P0442 Gongora claviodora Costa Rica Serrano 125 20.07.2007<br />

P0227 Gongora gale<strong>at</strong>a Ecuador 17.05.2006<br />

P03 13 Gongora gale<strong>at</strong>a Mexico 29.09.2006<br />

POOlS Gongora leucochila Costa Rica 19.01.2004<br />

P0226 Gongora trunc<strong>at</strong>a Venezuela 17.05.2006<br />

P0316 Gongora unicolor Costa Rica 29.09.2006<br />

P03 11 Gongora unicolor Costa Rica Pupulin 4954 29.09.2006<br />

P0229 Govenia ciliilabia Costa Rica PL/pulin 621 7 06.06.2006<br />

P0332 Gramm<strong>at</strong>ophyllum multiflorum Philippines 26.10.2006<br />

POO17 Gramm<strong>at</strong>ophyliL/m scriptum Philippines 12.05.2004<br />

P0493 Guarian<strong>the</strong> skinneri Costa Rica 05.02.2008<br />

POO14 Haraella retrocalla Taiwan 30.10.2002<br />

POJ05 Heterotaxis sessilis Costa Rica Clark 209 J9.05.2005<br />

P0324 Houlletia tigrina Costa Rica Bogar[n 2480 03.10.2006

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T ABLE 1. Continued.<br />

PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 75<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

P0051 HuntLeya burtii Costa Rica 13.05.2005<br />

P0402 Huntleya burtii Costa Rica 19.07.2007<br />

POO92 Hygroehilus parishii Philippines 13 .05.2005<br />

P0200 Kefersteinia Costa Rica Bogarfn 806 09.05.2006<br />

P0197 Kefersteinia andreettae Ecuador 02.05.2006<br />

P0395 Kefersteinia aurieuL<strong>at</strong>a Panama DressLer 6831 12.03.2006<br />

P0417 Kefersteinia eostarieensis Costa Rica Bogarfn 761 19.07.2007<br />

P0056 Kefersteinia eostarieensis Costa Rica Pupulin 5029 13.05.2005<br />

P0052 Kefersteinia exeentriea Costa Rica BLanco 1053 13.05.2005<br />

P0426 Kefersteinia Lamin<strong>at</strong>a Ecuador Pupulin 655/ 19.07.2007<br />

P0385 Kefersteinia orbieuLaris Costa Rica HoLst 8773 07.03.2006<br />

P0155 Kefersteinia orbieuLaris Costa Rica Pupulin s.n. 13.05.2005<br />

P0057 Kefersteinia orbieuLaris Costa Rica PupuLin s.n. 13.05.2005<br />

P0055 Kefersteinia parviLabris Costa Rica Bogarfn 143 13.05.2005<br />

POl98 Kefersteinia pellita Ecuador Pupulin 5998 02.05.2006<br />

P023 1 Kefersteinia retanae Costa Rica Cambronero s.n. 06.06.2006<br />

P0427 Kefersteinia taurina Ecuador Pupulin 6550 19.07.2007<br />

P0193 Kefersteinia taurina Ecuador Pupulin 5892 17.03.2006<br />

POO04 Kefersteinia toLùnensis Colombia Pupulin 4205 30.10.2002<br />

P0420 Kefersteinia toLimensis Colombia Pupulin 4205 19.07.2007<br />

P0054 Kefersteinia tolimensis Colombia Pupulin 4205 13.05.2005<br />

P0053 Kefersteinia werekLei Costa Rica PupuLin 3039 13.05.2005<br />

P044 1 Koellensteinia graminea Ecuador PupuLin 6517 20.07.2007<br />

P0165 LeoehiLus tricuspid<strong>at</strong>us Costa Rica PupuLin 5648 07.07.2005<br />

P0128 LeucohyLe subuL<strong>at</strong>a Nicaragua PupuLin 4301 09.06.2005<br />

P0090 Loekhartia amoena Costa Rica PupuLin 4515 13.05.2005<br />

P0091 Loekhartia pittieri Costa Rica BLanco 2653 13.05.2005<br />

P0184 Lyeasle Costa Rica 22. 12.2005<br />

POl 87 Lyeaste Costa Rica 29.12.2005<br />

P0217 Lyeaste Costa Rica 03.05.2006<br />

P0393 Lycaste Costa Rica 07.03.2006<br />

P0454 Lyeaste Costa Rica 20.07.2007<br />

P0462 Lyeaste Costa Rica Pupulin 2780 20.07.2007<br />

P0444 Lyeaste deppei México- 20.07.2007<br />

Nicaragua<br />

P0230 Lycaste deppei México- 06.06.2006<br />

Nicaragua<br />

P0460 Lyeaste Leueantha Costa Rica 20.07.2007<br />

P0437 Lycaste leucantha Costa Rica Bogarfn 2070 20.07.2007<br />

P0024 Lyeaste Leucantha Costa Rica 03.10.2002<br />

P0334 Lycaste x Michelii 26.10.2006<br />

P0476 Lyeaste sehilleriana Panama 19.08.2007<br />

P0201 Lycaste tricoLor Costa Rica 09.05.2006<br />

P0383 Lyeaste tricoLor Costa Rica 07.03.2006<br />

P0391 Masdevallia reiehenbachiana Costa Rica Vega s.n. 07.03.2006<br />

P0129 Maxillaria Costa Rica 22.1 2.2005<br />

P0180 Max illaria Costa Rica 22.12.2005<br />

P0357 Maxillaria Costa Rica 14.11.2006<br />

P0361 Maxillaria Costa Rica 14.11.2006<br />

P0364 Maxillaria Costa Rica BLanco 2808 14.1 1.2006<br />

P0235 Maxillaria Costa Rica BLanco 2808 06.06.2006<br />

P0178 Maxillaria Costa Rica G6mez 40 22.1 2.2005<br />

P0240 Maxillaria Costa Rica Pupulin 3023 13.06.2006<br />

P0028 Maxillaria Costa Rica Whitten 2045 05.05.2006<br />

PO 107 Max illaria aeostae Costa Rica BLanco 941 19.05.2005<br />

PO 103 Maxillaria aLba Costa Rica PupuLin 2850 19.05.2005<br />

POIOI Maxillaria angustissima Costa Rica 19.05.2005<br />

P0167 Maxillaria araehnitiflora Costa Rica Bogarfn 344 22.12.2005<br />

P0262 Maxillaria araehnitifiora Costa Rica BLanco 2262 30.06.2006<br />

P0095 Maxillaria <strong>at</strong>woodiana Costa Rica 18.05.2005<br />

P0234 Maxillaria <strong>at</strong>woodiana Costa Rica 06.06.2006<br />

POI04 Maxillaria braehybulbon Costa Rica Bogarfn 874 19.05.2005

---

76 SELBYANA Volume 29(1) 2008<br />

TABLE 1. Continued.<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

P0249 Maxillaria brunnea Costa Rica Whitten 2028 16.06.2006<br />

P0189 Maxillaria caespitifica Costa Rica Blanco 1076 22.12.2005<br />

P0268 Maxillaria campanul<strong>at</strong>a Costa Rica Bogarfn 1801 12.07.2006<br />

PO 179 Maxillaria campanul<strong>at</strong>a Costa Rica Comez 48 22.12.2005<br />

P0259 Maxillaria confusa Costa Rica Bogarfn 1324 30.06.2006<br />

P0236 Maxillaria con/usa Costa Rica Whitten 2030 06.06.2006<br />

P0365 Maxillaria con/usa Costa Rica Whitten 2030 14.1 1.2006<br />

P0181 Maxillaria endresii Costa Rica Whitten 2089 22.12.2005<br />

P0097 Maxillaria endresii Costa Rica 18.05.2005<br />

P0264 Maxillaria haberi Costa Rica Bogar{n 1868 05.07.2006<br />

P0093 Maxillaria hedwigae Costa Rica Pupulin J 859 18.05.2005<br />

P0176 Maxillaria inaudita Costa Rica Comez S.n. 22.12.2005<br />

P0106 Maxillaria lueri Costa Rica 19.05.2005<br />

POO09 Maxillaria luteoalba Colombia 06.11.2002<br />

P0244 Maxillaria meridensis Costa Rica Comez 19 13.06.2006<br />

P0099 Maxillaria moralesii Costa Rica 18.05.2005<br />

P0248 Maxillaria moralesii Costa Rica Pupulin 4682 16.06.2006<br />

PO 102 Maxillaria nasuta Costa Rica Pupulin 3247 19.05.2005<br />

PO 108 Maxillaria pseudoneglecta Costa Rica 19.05.2005<br />

POI77 Maxillaria punctostri<strong>at</strong>a Costa Rica Comez 1 22.12.2005<br />

P0094 Maxillaria ramonensis Costa Rica 18.05.2005<br />

P0242 Max!llaria reichenheimiana Costa Rica Blanco 2107 13.06.2006<br />

POO96 Maxillaria reichenheimiana Costa Rica 18.05.2005<br />

POOlO Maxillaria ringens Costa Rica 21.1 1.2002<br />

P0366 Maxillaria ringens Costa Rica Comez 24 14.11.2006<br />

POlOO Maxillaria ringens Costa Rica Comez 24 18.05.2005<br />

P0098 Maxillaria rodrigueziana Costa Rica 18.05.2005<br />

P0232 Maxillaria rodrigueziana Costa Rica 06.06.2006<br />

P0239 Maxillaria sanguinea Costa Rica 07.06.2006<br />

PO 109 Maxillaria schlechteriana Costa Rica Whitten 2/81 19.05.2005<br />

POl57 Maxillaria scorpioidea Costa Rica 22.12.2005<br />

P0321 Maxillaria serrul<strong>at</strong>a Costa Rica Bogarfn 1135 03.10.2006<br />

P024 1 Maxillaria tonduzii Costa Rica Blanco 3179 13.06.2006<br />

P0243 Maxillaria wercklei Costa Rica Blanco 3179 13.06.2006<br />

P0371 Mesospinidium horichii Costa Rica Pupulin 5357 25.01.2006<br />

P0369 Mesospinidium panamense Panama Dressler s.n. 25.01.2006<br />

P0370 Mesospinidium warszewiczii Costa Rica 25.01.2006<br />

P0470 Micochilus biment<strong>at</strong>us Costa Rica Pupulin 6690 13.08.2007<br />

P0074 Miltonia regnellii Brazil 13.05.2005<br />

P0080 Miltoniopsis phalaenopsis Colombia 13.05.2005<br />

P0078 Miltoniopsis roezlii Colombia 13.05.2005<br />

P0079 Miltoniopsis vexillaria Colombia 13.05.2005<br />

POO77 Miltoniopsis warscewiczii Costa Rica 13.05.2005<br />

P0382 Mormodes colossa Costa Rica Blanco 2691 07.03.2006<br />

P0443 Mormodes fractiflexa Costa Rica Bogarfn 2705 20.07.2007<br />

P0223 Mormodes horichii Costa Rica. Bogarfn 2561 12.05.2006<br />

P0023 Mormodes <strong>at</strong>ropurpurea Costa Rica 04.03.2005<br />

P0190 Mormodes colossa Costa Rica Blanco 2192 22.12.2005<br />

P0081 Mormolyca ringens Costa Rica 13.05.2005<br />

P0215 Oncidium Costa. Rica 03.05.2006<br />

P0214 Oncidium Costa Rica Comez 18 03.05.2006<br />

P0034 Oncidium coquianum Costa Rica Pupulin 3438 19.04.2006<br />

P0375 Oncidium dichrom<strong>at</strong>icum Costa Rica 28.02.2006<br />

P0434 Oncidium dichrom<strong>at</strong>icum Costa. Rica Pupulin 2495 19.07.2007<br />

P0373 Oncidium ens<strong>at</strong>um Ecua.dor Pupulin 6421 05.02.2006<br />

P0372 Oncidium fusc<strong>at</strong>um Nicaragua Pupulin 6420 05.02.2006<br />

P0374 Oncidium hyphaem<strong>at</strong>icum Nicaragua Pupulin 6422 05.02.2006<br />

P0432 Oncidium hyphaem<strong>at</strong>icum N icaragua. Bogarfn 3793 19.07.2007<br />

P0431 Oncidium kotzchianum Costa Rica Pupulin 5455 19.07.2007<br />

P0331 Oncidium luteum Costa. Rica. Pupulin 6197 26.10.2006<br />

P0182 Oncidium luteum Costa Rica Whitten 2192 22.12.2005<br />

P01I3 Oncidium oliganthum Gu<strong>at</strong>emala Pupulin 3378 24.05.2003

---

T ABLE 1. Continued.<br />

PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 77<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

POIIl Oncidium parviflorum Costa Rica 24.05.2003<br />

P0458 Oncidium parviflorum Costa Rica Bogarfn 2468 20.07.2007<br />

POl83 Oncidium schroederianum Costa Rica 22.1 2.2005<br />

P0440 Oncidium schroederianum Costa Rica Pupulin 6254 20.07.2007<br />

P0418 Oncidium sp.nov. Costa Rica Karremans 1650 19.07.2007<br />

P0333 Oncidium stenobulbon Costa Rica Bogar(n 1065 26.10.2006<br />

POI14 Oncidium storkii Costa Rica 24.05.2003<br />

P0250 Osmoglossum egertonii Costa Rica 16.06.2006<br />

P0266 Otoglossum chiriquense Costa Rica Pupulin 6235 12.07.2006<br />

P0082 Otoglossum globuliferum Costa Rica Pupulin 5296 13.05.2005<br />

P0273 Paphinia subclausa Costa Rica Dressler 6590 01.09.2006<br />

P0455 Paphinia subclausa Costa Rica Dressler 6590 20.07.2007<br />

POl61 Paphinia subclausa Costa Rica Pupulin 4665 07.07.2005<br />

P0446 Pesc<strong>at</strong>orea cerina Costa Rfca Blanco 2642 20.07.2007<br />

P0389 Pesc<strong>at</strong>orea cerina Costa Rica Pupulin 2993 07.03.2006<br />

P0068 Pesc<strong>at</strong>orea cerina Costa Rica Pupulin 2097 17.07.2005<br />

P0061 Pesc<strong>at</strong>orea cerina Costa Rica 13.05.2005<br />

POO07 Pesc<strong>at</strong>orea cerina Costa Rica 06.08.2003<br />

P0062 Pesc<strong>at</strong>orea dayana Colombia 13.05.2005<br />

POI92 Phalaenopsis amabilis Asia Pupulin 5713 17.03 .2006<br />

POO02 Phalaenopsis schilleriana Philippines 12.05.2004<br />

P0173 Phalaenopsis amabilis lndonesia Pupulin 5713 17.07.2005<br />

P0397 Phalaenopsis luedemanniana Philippines 28.03.2006<br />

POO18 Phalaenopsis violacea Malasya 22.10.2005<br />

P0472 Polycicnis barb<strong>at</strong>a Costa Rica Bogarfn 1898 19.08.2007<br />

P0022 Polycycnis barb<strong>at</strong>a Costa Rica 04.03.2005<br />

P0265 Polycycnis barb<strong>at</strong>a Costa Rica Bogarfn 1898 12.07.2006<br />

P0314 Polycycnis barb<strong>at</strong>a Costa Rica Bogarfn 1845 29.09.2006<br />

P0063 Pros<strong>the</strong>chea brassavolae Costa Rica 22.12.2005<br />

P0392 Pros<strong>the</strong>chea chacaoensis Costa Rica 07.03.2006<br />

POl34 Pros<strong>the</strong>chea neglecta Costa Rica 10.06.2005<br />

POl62 Pros<strong>the</strong>chea chacaoensis Costa Rica 07.07.2005<br />

P0133 Pros<strong>the</strong>chea vitellina Mexico 10.06.2005<br />

P0116 Rodriguezia venusta Ecuador Pupulin 5281 08.06.2005<br />

P0158 Rodriguezia venusta Ecuador 16.05.2005<br />

P0270 Rossioglossum schlieperianum Costa Rica Bogarfn 2851 18.07.2006<br />

P0083 Rossioglossum schlieperianum Costa Rica 13 .05.2005<br />

POll2 Rudolfiella aurantiaca Ecuador 24.05 .2003<br />

P0135 Schomburgkia lueddemannii Costa Rica 13.06.2005<br />

POOll Sedirea japonica Japan 12.05 .2004<br />

P0463 Sievekingia fimbri<strong>at</strong>a Costa Rica Bogarfn 2813 20.07.2007<br />

P0445 Sievekingia fimbri<strong>at</strong>a Costa Rica 20.07.2007<br />

P0136 Sievekingia fimbri<strong>at</strong>a Costa Rica 13.06.2005<br />

P0209 Sievekingia fimbri<strong>at</strong>a Costa Rica 03.05.2006<br />

P0433 Sievekingia suavis Costa Rica Bogarfn 2575 19.07.2007<br />

P0438 Sigm<strong>at</strong>ostalix cardioglossa Costa Rica Pupulin 3499 20.07.2007<br />

P0233 Sigm<strong>at</strong>ostallx dulcineae Costa Rica Bogar(n 2079. 06.06.2006<br />

P0356 Sobralia Costa Rica Aguilar 13 14.11.2006<br />

P0338 Sobralia Costa Rica Bogarfn 913 14.11.2006<br />

P0306 Sobralia Costa Rica Dressler 6723 13.09.2006<br />

P0337 Sobralia Costa, Rica Dressler 6621 14.11.2006<br />

P0363 Sobralia Costa Rica Karremans 661 14.11.2006<br />

P0260 Sobralia amabilis Costa Rica Dressler 6713 30.06.2006<br />

P0367 Sobralia andreae Colombia 14.11.2006<br />

P025 1 Sobralia hel/eri Costa Rica Dressler 6539 21.06.2006<br />

P0492 Sobralia luteola Costa Rica PupiAlin 2629 05 .02.2008<br />

P0343 Sobralia macra Costa, Rica 14.11.2006<br />

P0259 Sobralia macrophyl/a Costa Rica Casteifranco 155 30.06.2006<br />

P0256 Sobralia quin<strong>at</strong>a Costa, Rica Pupulin 3664 30.06.2006<br />

P0160 Sobralia leuçoxantha Costa Rica Pupulin 4184 07.07.2005<br />

P0448 Stanhopea Costa Rica 20.07.2007<br />

P0263 Stanhopea Costa Rica 05.07.2006

---

78 SELBYANA Volume 29(1) 2008<br />

TABLE,1. Continued.<br />

No. Genus/species Country Collector D<strong>at</strong>e<br />

P0267 Stanhopea Costa Rica 12.07.2006<br />

P0294 Stanhopea Costa Rica 05.09.2006<br />

P0430 Stanhopea Costa Rica 19.07.2007<br />

P0452 Stanhopea Costa Rica 20.07.2007<br />

P0277 Stanhopea Costa Rica Bogarfn 1639 01.09.2006<br />

P0387 Stmihopea Costa Rica Bogarfn 757 07.03.2006<br />

P0276 Stanhopea Costa Rica Pupulin 2960 01.09.2006<br />

P0291 Stanhopea Costa Rica Pupulin 2960 05.09.2006<br />

P0280 Stanhopea Costa Rica Rojas 112 05.09.2006<br />

P0285 Stanhopea Ecuador Pupulin 6279 05.09.2006<br />

P0292 Stanhopea Nicaragua Karremans 557 05.09.2006<br />

P0210 Stanhopea Venezuela 03.05.2006<br />

P0257 Stanhopea Venezuela 30.06.2006<br />

P0247 Stanhopea cfcostaricensis Costa Rica 13.06.2006<br />

P0290 Stanhopea cirrh<strong>at</strong>a Costa Rica Bogarfn 1021 05.09.2006<br />

P0204 Stanhopea cirrh<strong>at</strong>a Costa Rica Bogarfn 1021 03.05.2006<br />

P0203 Stanhopea cirrh<strong>at</strong>a Costa Rica Castelfranco 153 03.05.2006<br />

P0295 Stanhopea cirrh<strong>at</strong>a Costa Rica Pupulin 2843 05.09.2006<br />

P0315 Stanhopea cirrh<strong>at</strong>a Costa Rica Pup ulin 5827 29.09.2006<br />

P0485 Stanhopea confusa Costa Rica Flores S.n. 02.10.2007<br />

P0466 Stanhopea costaricensis Costa Rica 20.07.2007<br />

P0225 Stanhopea costaricensis Costa Rica 16.05.2006<br />

P045 1 Stanhopea costaricensis Costa Rica 20.07.2007<br />

P0457 Stanhopea costaricensis Costa Rica 20.07.2007<br />

P0474 Stanhopea costaricensis Costa Rica 19.08.2007<br />

P0205 Stanhopea costaricensis Costa R ica 03.05.2006<br />

P0368 Stanhopea costaricensis x cirrh<strong>at</strong>a Costa Rica Rojas 113 26.11.2006<br />

P0254 Stanhopea ecornuta Costa Rica Rojas 110 2 1.06.2006<br />

P0283 Stanhopea ecornuta Costa Rica 05 .09.2006<br />

P0288 Stanhopea ecornuta Costa R ica 05.09.2006<br />

P0289 Stanhopea ecornuta Costa Rica 05.09.2006<br />

P0137 Stanhopea ecornuta Costa Rica 13.06.2005<br />

P0305 Stanhopea ecornuta Costa Rica 13.09.2006<br />

P0471 Stanhopea ecornuta Costa Rica 19.08.2007<br />

P0202 Stanhopea x fowlieana Costa Rica 03.05.2006<br />

P0253 Stanhopea x fowlieana Costa Rica 21.06.2006<br />

P0252 Stanhopea gibbosa Costa Rica 21.06.2006<br />

P0459 Stanhopea gibbosa Costa Rica 20.07.2007<br />

P01 40 Stanhopea gibbosa Costa Rica 13.06.2005<br />

P0449 Stanhopea grandiflora 20.07.2007<br />

P0330 Stanhopea ocul<strong>at</strong>a Mexico Rojas 134 26.10.2006<br />

P0345 Stanhopea ocul<strong>at</strong>a Mexico Rojas 135 14.11.2006<br />

P0353 Stanhopea pulla Costa Rica Rojas 136 14. 11.2006<br />

P0206 Stanhopea pulla Costa Rica 03.05.2006<br />

P0208 Stanhopea pulla Costa Rica 03.05.2006<br />

P0020 Stanhopea pulla Costa Rica 21.06.2003<br />

P0344 Stanhopea wardii Costa Rica Bogarfn 630 14.11.2006<br />

P0328 Stanhopea wardii Costa Rica Dressler 6564 17.10.2006<br />

P0335 Stanhopea wardii Costa Rica G6mez 51 26.10.2006<br />

P0296 Stanhopea wardii Costa Rica Pupulin 4930 05 .09.2006<br />

P0484 Stanhopea wardii Costa Rica Pupulin 4390 02.10.2007<br />

POO16 Stanhopea wardii Costa Rica Pupulin 11 79 20.07.2004<br />

P0146 Stanhopea wardii Costa Rica Pupulin 4930 13.06.2005<br />

P0147 Stanhopea wardii Costa Rica Pupulin 1179 13.06.2005<br />

P0207 Stanhopea wardii Costa Rica 03.05.2006<br />

P0482 Stanhopea wardii Costa Rica 28.09.2007<br />

P0483 Stanhopea wardii Costa Rica 28.09.2007<br />

P0487 Stanhopea wardii Costa Rica 25.10.2007<br />

POO19 Stanhopea wardii Costa Rica 05.08.2003<br />

P0138 Stanhopea wardii Costa Rica 13.06.2005<br />

P0139 Stanhopea wardii Costa Rica 13.06.2005<br />

P014 1 Stanhopea wardii Costa R ica 13.06.2005<br />

P0142 Stanhopea wardii Costa Rica 13.06.2005

---

TABLE 1. Continued.<br />

PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 79<br />

No. Genus/species Country Collector Dale<br />

P0144 Stanhopea wardii Costa Rica 13.06.2005<br />

P0145 Stanhopea wardii Costa Rica 13.06.2005<br />

P0293 Stanhopea warscewicziana Costa Rica 05.09.2006<br />

P0297 Stanhopea warscewicziana Costa Rica 06.09.2006<br />

P0304 Stanhopea warscewicziana Costa Rica 13.09.2006<br />

P0327 Stanhopea warscewicziana Costa Rica 10.10.2006<br />

P0143 Stanhopea warscewicziana Costa Rica 13.06.2005<br />

P0269 Stenia Ecuador Pupulin 6239 18.07.2006<br />

P0348 Stenia calceolaris Ecuador Pupulin 6332 14.1 1.2006<br />

P0350 Stenia dodsoniana Ecuador Pupulin 6340 14.11.2006<br />

P0279 Stenia gutt<strong>at</strong>a Ecuador Pupulin 5937 05.09.2006<br />

P0318 Stenia pallida Ecuador Pupulin 6286 29.09.2006<br />

P0415 Stenia pallida Ecuador 19.07 .2007<br />

P0191 Stenia stenioides Ecuador Pupulin 5893 17.03.2006<br />

P0407 Ste nia stenioides Ecuador Pupulin 5893 19.07.2007<br />

P0199 Stenotyla <strong>lankester</strong>iana Costa Rica 02.05.2006<br />

POl52 Stenotyla <strong>lankester</strong>iana Costa Rica 19.04.2006<br />

P0066 Stenotyla lendyana Gu<strong>at</strong>emala Hubein s.n. 13.05.2005<br />

P0065 Stenotyla lendyana Gu<strong>at</strong>emala 13.05.2005<br />

P0421 Stenotyla lendyana Gu<strong>at</strong>emala 19.07.2007<br />

P0245 Stenotyla panamensis Panama Pupulin 5868 13.06.2006<br />

P0477 Stenotyla panamensis Panama Pupulin 5869 19.08.2007<br />

P0064 Stenotyla picta Costa Rica Pupulin 3031 13.05.2005<br />

P0422 Stenotyla picta Costa Rica 19.07.2007<br />

P0423 Stenotyla picta Costa Rica 19.07.2007<br />

P0384 Stenotyla picta Costa Rica Dressler s.n. 07.03.2006<br />

P0435 Stenotyla picta Panama Bogarfn 2960 19.07.2007<br />

P0298 TeLipogon biolleyi Costa Rica Karremans 1317 06.09.2006<br />

P0237 Telipogon costaricensis Costa Rica Rojas 7148 06.06.2006<br />

P0238 Telipogon monticola Costa Rica Rojas 7157 06.06.2006<br />

P0148 TeLipogon ampliflorus Costa Rica 13.06.2005<br />

P0073 Ticoglossum krameri Costa Rica 13.05.2005<br />

P0070 Ticoglossum oerstedii Costa Rica 13.05.2005<br />

P0220 Tolumnia triquetra Jamaica 03.05.2006<br />

P0323 Trevoria zalhbrukneriana Costa Rica Bogarfn 1900 03.10.2006<br />

P0211 Trevoria zalhbrukneriana Costa Rica G6mez 39 03.05.2006<br />

P0085 Trichocentrum capistr<strong>at</strong>um Costa Rica Pupulin 2075 13.05.2005<br />

P0084 Trichocentrum cymbiglossum Costa Rica PupuLin 4754 13.05.2005<br />

P0377 Trichocentrum pfavii Costa Rica Pupulin 5826 05.03.2006<br />

P0360 Trichoglottis Philippines 14.11.2006<br />

P0424 Trichopilia Colombia Dressler 6807 19.07.2007<br />

P0349 TrichopiLia Colombia PupuLin 6333 14.11.2006<br />

P0490 Trichopilia Costa Rica Pupulin 641 8 14.01.2008<br />

P0399 Trichopilia eneidae Panama 28.03.2006<br />

P0491 Trichopilia fragrans Colombia 14.01 .2008<br />

P0486 Trichopilia fragrans Colombia Pupulin 6341 25.10.2007<br />

P0351 Trichopilia aff. fragrans Colombia Pupulin 6341 14.11 .2006<br />

POl69 Trichopilia galeottiana Costa Rica Bogarfn 145 07.07.2005<br />

P0400 Trichopilia hennisiana Colombia 28.03.2006<br />

POl51 Trichopilia margin<strong>at</strong>a Costa Rica 14.06.2005<br />

P0076 Trichopilia margin<strong>at</strong>a Costa Rica Pupulin 4382 13.05.2005<br />

P0175 Trichopilia margin<strong>at</strong>a Costa Rica Pupulin 4433 17.07.2005<br />

P0309 Trichopilia nobilis Colombia 20.09.2006<br />

P0453 Trichopilia olmosii Panama Bogarfn 2948 20.07.2007<br />

P0075 Trichopilia punicea Costa Rica Pupulin 5213 13.05.2005<br />

poon Trichopilia punicea Costa Rica 13.05.2005<br />

P0320 Trichopilia turrialbae Costa Rica 03. l0.2006<br />

P021 2 Vanda coerulea Philippines 03.05.2006<br />

P0213 Vanda lamell<strong>at</strong>a Philippines 03.05.2006<br />

P04 4 Warczewiczella discolor Costa Rica 19.07.2007<br />

P0059 Warscewiczella discolor Costa Rica 13.05.2005<br />

P0058 Warczewiczella ionoleuca Ecuador PupuLin 5235 13.05.2005<br />

P0401 Warczewiczella ionoleuca Ecuador PupuLin 5235 19.07.2007

---

82 SELBYANA Volume 29(1) 2008<br />

F IGURE 6. Represent<strong>at</strong>ive <strong>pollinaria</strong> of <strong>the</strong> Zygopetalinae. 1-8: Dichaea. 1-D. truila (Pupulin 5130); 2-<br />

D. calycul<strong>at</strong>a (Pupulin 6923); 3-D. globosa (Pupulin 5319); 4-D. globosa (JBL-09172); 5-Dichaea sp.<br />

(Pupulin 6891); 6-D. morrisii (Whitten 2171); 7- D. poiciilantha (Karremans 1140); 8- D. obov<strong>at</strong>ipetala<br />

(JBL-08203). 9-13: Daiotyla. 9-D. xanthina (Pupulin 5891); lü,lI- D. albicans (Pupulin 4372); 12, 13-D.<br />

crassa (Pupulin 4257). 14- 16: Chondroscaphe. 14-C. amabilis (Pupulin 6553); 15- D. escobariana (Pupulin<br />

6289); 16-D. flaveola (Pupulin 6474). 17-27: Kefersteinia. 17- K. costaricensis (Bogarfn 761); 18-K.<br />

lamin<strong>at</strong>a (Pupulin 6551); 19- K. o rbicularis (Pupulin 2068); 20-K. retanae (JBL-011 13);

---

PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 83<br />

(Continued) 2 1-K. wercklei (JBL-03465); 22- K. excentrica (Pupulin 5225); 23-K. excentrica (JBL-00469);<br />

24-K. parvilabris (JBL-05186); 25-K. tolimensis (Pupulin 4205); 26-K. mystacina (Pupulin 6559); 27- K.<br />

taurina (Pupulin 6550). 28-32: Benzingia. 28- B. reichenbachiana (Bogarîn 1890); 29, 30-B. reichenbachiana<br />

(JBL-07994); 31-B. cornuta (Pupulin 5879); 32- B. caud<strong>at</strong>a (Pupulin 6544). 33: xBensteinia ramonensis<br />

(Bogarîn 1923). 34: B<strong>at</strong>emannia leferenzii (Pupulin 6224). 35: Zygopetalinae gen. ined. (Pupulin 6605). 36-<br />

41 : Chaubardiella. 36-C pubescens (Pupulin 6240); 37-C aff. pubescens (Pupulin 6566); 38-C subquadr<strong>at</strong>a<br />

(Pupulin 5716); 39, 40-C pacuarensis (JBL-0111 7, JBL-05422); 4t-C aff. subquadr<strong>at</strong>a (Pupulin 6564).<br />

42- 48: Stenotyla. 42, 43- S. <strong>lankester</strong>iana (JBL-OJ651); 44, 45-S. picta (JBL-OJ970, JBL-01905); 46, 47-S.<br />

lendyana (JBL-02053, JBL-0896); 48-S. panamensis (Pupulin 5868). 49- 56: Stenia. 49- S. gutt<strong>at</strong>a (Pupulin<br />

6567); 50-S. dodsoniana (Pupulin 6340); 51-S. stenioides (Pupulin 5893); 52-S. angustilabia (Pupulin<br />

6570); 53-S. calceolaris (Pupulin 6332); 54, 55-S. pallida (Pupulin 6286; Pupulin 6548); 56-S. lillianae<br />

(Pupulin 6557). 57-62: Warczewiczella. 57, 58- W. discolor (Pupulin 3939; JBL-04925); 59- W. margin<strong>at</strong>a<br />

(Pupulin 6516); 60, 61-W. ionoleuca (Pupulin 5235; JBL-00873);62- W. pal<strong>at</strong>ina (Pupulin 6909). 63-65:<br />

Cochlean<strong>the</strong>s. 63, 64-C arom<strong>at</strong>ica (JBL-02932; JBL-07583); 65-C flabelliformis (Pupulin 6621). 66: Zygosepalum<br />

lindeniae (Pupulin 6643). 67: Ixyophora viridisepala (Pupulin 6554). 68: "Chondrorhycnha " velastiguii<br />

(Pupulin 6622). 69- 72: Pesc<strong>at</strong>orea. 69, 70, 71- P. cerina (JBL-09435; JBL-09430; JBL-06217); 72- P.<br />

dayana (JBL-03943). 73: Zygopetalum intermedium (Pupulin s.n.). 74: Galeottia grandiflora (JBL-J4627). 75-<br />

78: Huntleya. 75, 76-H. burtii (JBL-14492; Pupulin 3829); 77-H. citrina (Pupulin 6645); 78-H. lucida<br />

(Pupulin 6642). 79-80: Euryblema andreae (Pupulin 6282). Scale bar = 5 mm.

---

84 SELBYANA Volume 29(1) 2008<br />

FIGURE 7. Pollinaria of Stanhopea species. 1- 16: S. wardii. I- Dressler 6564; 2 - JBL-04360; 3 - JBL-<br />

06666; 4 - DB 630; 5 - JBL-07741; 6 - R . Gomez 51; 7 - JBL-06133 ; 8 - FP 1179; 9 - JBL-04363; 10 - FP<br />

4390; 11 - JBL-09981 ; 12 - JBL-07743; 13 - FP 4930; 14 - JBL-06126; 15 - JBL-0615J ; 16 - FP 4930. 17-<br />

21: S. warscewicziana.17 - JBL-14610; 18 - JBL-08955; 19 - JBL-06148; 20 - JBL-14605; 21 - JBL-07747. 22-<br />

28: S. costaricensis. 22 - JBL-05094; 23 - JBL-05659; 24 - JBL-07726; 25 - JBL-05094; 26 - JBL-05066; 27 -<br />

JBL-09988; 28 - JBL-05069. 29-35: S. ecomuta. 29 - JBL-09867; 30 - JBL-1l353; 31 - JBL-12602; 32 - JBL­<br />

JJ696; 33 - JBL-06132; 34 - Rojas 110; 35 - JBL-11698. 36- 39: S. pulla. 36 - JBL-05657; 37

---

PUPULIN & KARREMANS: POLLINARIA AT LANKESTER 85<br />

TABLE 2. Polli naria included in <strong>the</strong> <strong>collection</strong>, arranged<br />

system<strong>at</strong>ically (phylogentic c lassific<strong>at</strong>ion<br />

according to Chase et al. 2003). Numbers in<br />

brackets refer to <strong>the</strong> number of <strong>pollinaria</strong> for each<br />

species or morphospecies.<br />

Orchidoideae<br />

Cranichideae<br />

Goodyerinae: Erythrodes tridax, Microchilus biment<strong>at</strong>us<br />

Spiranthinae: Coccineorchis standleyi<br />

Epidendroideae<br />

Sobralieae: Sobralia amabilis, S. andreae, S. helleri,<br />

S. luteola, S. mac ra, S. macrophylla, S.<br />

quin<strong>at</strong>a, S. leucoxantha, Sobralia ssp. (5)<br />

Calypsoeae: Govenia ciliilabia<br />

Epidendreae<br />

Pleurothallidinae: Masdevallia reichenbachiana<br />

Laeliinae : Arpophyllum giganteum, Barkeria lindleyana,<br />

Brassavola nodosa, C<strong>at</strong>tleya inlermedia,<br />

Encyclia ambigua, E. cordigera, E. cer<strong>at</strong>istes,<br />

Epidendrum centradenia, E.<br />

odontochilum, E. schumannianum, E. wallisii,<br />

Epidendrum spp. (3), Guarian<strong>the</strong> skinneri,<br />

Pros<strong>the</strong>chea brassavolae, P. chacaoensis (2),<br />

P. neglecta, P. vitellina, Schomburgkia lueddemannii<br />

Coeliinae [?): Coelia bella<br />

Cymbidieae<br />

C<strong>at</strong>asetinae: C<strong>at</strong>asetum macul<strong>at</strong>um (3), Cycnoches<br />

amparoanum, Cyrtopodium macrobulbon,<br />

Dressleria dilecta, Galeandra lacustris, Mormodes<br />

<strong>at</strong>ropurpurea, M. colossa (2), M. fractiflexa,<br />

M. horichii<br />

Cymbidiinae: Cymbidium iridioides (2), Gramm<strong>at</strong>ophyllum<br />

multiflorum, G. scriptum<br />

Eriopsidinae: Eriopsis wercklei (2)<br />

Oncidiinae: Ada aurantiaca (2), A. chlorops, Aspasia<br />

epidendroides, Brassia arcuigera (6), B. caud<strong>at</strong>a<br />

(4), B. gireoudiana (2), B. suavissima, B.<br />

verrucosa, Comparettia falc<strong>at</strong>a, Erycina cristagalli,<br />

Leochilus tricuspid<strong>at</strong>us, Lockhartia amoena,<br />

L. pittieri, Mesospinidium horichii, M. panamense,<br />

M. warszewiczii, Miltonia regnellii, Miltoniopsis<br />

phalaenopsis, M. roezlii, M. vexillaria, M.<br />

warscewiczii, Oncidium coquianum, O. dichrom<strong>at</strong>icum,<br />

O. dichrom<strong>at</strong>icwn, 0. ens<strong>at</strong>um, O. fusc<strong>at</strong>um,<br />

O. hyphaem<strong>at</strong>icum (2), 0. klotzchianum, O.<br />

luteum (2), O. oliganthum, O. parviflorum (2), O.<br />

parviflorum, O. schroederianum (2), 0. stenobulbon,<br />

O. storkii, Oncidium spp. (3), Osnwglossum<br />

egertonii, Otoglossum chiriquense, 0. globuliferum,<br />

Rodriguezia venusta (2), Rossioglossum<br />

schlieperianum (2), Sigm<strong>at</strong>ostalix dulcineae, S.<br />

cardioglossa, Telipogon ampliflorum, T. biolleyi,<br />

T. costaricensis, T. monticola, Ticoglossum<br />

krameri, T. oerstedii, Tolumnia triquelra, Trichocenlrum<br />

capistr<strong>at</strong>um, T. cymbiglossum, T. pfavii,<br />

f--<br />

TABLE 2. Continued.<br />

Trichopilia eneidae, T. fragans (2), T. aff. fragrans,<br />

T. galeottiana, T. hennisiana, T. margin<strong>at</strong>a<br />

(3), T. nobilis. T. olmosii, T. punicea (2), T. subu­<br />

L<strong>at</strong>a, T. turrialbae, Trichopilia spp. (3)<br />

Maxillariinae: Helerotaxis sessilis, Lycasle borucana<br />

(2), L. deppei (2), L. leucantha (3), L. michelii<br />

(lassioglossa X cochle<strong>at</strong>a), L. schilleriana, L. tricolor<br />

(2), Lycaste spp. (4), Maxillaria acostae, M .<br />

alba, M. angustissima, M . arachnitiflora (2), M.<br />

<strong>at</strong>woodiana (2), M. brachybulbon, M. cf. brunnea,<br />

M. caespitifica, M. campanul<strong>at</strong>a (2), M. confuse<br />

(3), M. endresii (2), M. haberi, M. hedwigae, M.<br />

inaudita, M. lue ri, M. luteoalba, M. meridensis, M.<br />

moralesii (2), M. nasuta, M. pseudoneglecta, M.<br />

punctostri<strong>at</strong>a, M . ramonensis, M. reichenheimiana<br />

(2), M. ringens (3), M. rodrigueziana (2), M. sanguinea,<br />

M. schlechteriana, M . scorpioidea, M. serrul<strong>at</strong>a,<br />

M. tonduzii, M. wercklei, Maxillaria spp.<br />

(9), Mormolyca ringens, Rudoljielia aurantiaca<br />

Stanhopeinae: Acineta densa (7), Acineta sp. (2),<br />

Gongora armeniaca (2), G. claviodora, G. gale<strong>at</strong>a<br />

(2), G. leucochila (2), G. trunc<strong>at</strong>a, G. unicolor (2),<br />

Gongora spp. (6), Houlletia tigrina, Paphinia subclausa<br />

(3), Polycicnis barb<strong>at</strong>a (4), Sievekingiajimbri<strong>at</strong>a<br />

(3), S. sua vis, Stanhopea cirrh<strong>at</strong>a (5), S.<br />

confusa, S. costaricensis (6), S. cf. costaricensis, S.<br />

costaricensis X cirrh<strong>at</strong>a, S. ecornuta (7), S. gibbosa<br />

(3), S. grandiflora, S. ocul<strong>at</strong>a (2), S. pulia (4),<br />

S. wardii (19), S. warscewicziana (5), S. X fowlieana<br />

(2), Stanhopea spp. (14), Trevoria zalhbrukneriana<br />

(2)<br />

Coeliopsidinae: Coeliopsis hyacinlhosma (2)<br />

Zygopetalinae: B<strong>at</strong>emania leferenzii, X Bensteinia<br />

ramonensis (2), Benzingia caud<strong>at</strong>a, B. cornuta, B.<br />

reichenbachiana (2), Chaubardia heteroclita,<br />

Chaubardiella pubescens, C. subquadr<strong>at</strong>a, C. pacuarensis<br />

(2), Chondroryncha thienii (2), Chondroscaphe<br />

amabilis, C. <strong>at</strong>rilinguis (2), C. bicolor (9),<br />

C. aff. bicolor (15), C. endresii (2), C. eburnea (2),<br />

C. embreei (2), C. escobariana, C. flaveola (3), C.<br />

cf. laevis (2), C. yamilethiae (5), Cochlean<strong>the</strong>s<br />

arom<strong>at</strong>ica (2), Daiotyla albicans (2), D. crassa<br />

(3), D. xanthina, Dichaea acroblephara (3), D.<br />

amparoana, D. cryptarrhena, D. elliptica (3), D.<br />

fragrantissima subsp. eburnea (2), D. globosa (3),<br />

D. morrisii (2), D. ovob<strong>at</strong>ipetala (2), D. panamensis<br />

(2), D. poicillantha (3), D. similis, D. cf. similis,<br />

D. trulla (2), Euryblema an<strong>at</strong>onum, E. andreae<br />

(2), Galeottia grandiflora (3), Huntleya burtii (2) ,<br />

Kefersteinia andreettae, K. auricul<strong>at</strong>a, K. costaricensis<br />

(2), K. excentrica, K. lamin<strong>at</strong>a, K. orbicu­<br />

Laris (2), K. pan1ilabris, K. pellita, K. retanae, K.<br />

taurina(2), K. tolimensis (3), K. wercklei, Kefersteinia<br />

spp. (2), Koellensteinia g raminea, Pesc<strong>at</strong>orea<br />

cerina (5), P. dayana, Stenic calceolaris, S.<br />

dodsoniana, S. gutt<strong>at</strong>a, Stenia pallida (2), Stenia<br />

stenoides (2), Stenotyla <strong>lankester</strong>iana (2), S. len-<br />

(Continued) - JBL-06155; 38 - Rojas 136; 39 - JBL-07727. 40-44: S. cirrh<strong>at</strong>a. 40 - DB 1021; 41- DB 1021;<br />

42 - FP 2843; 43 - FP 583; 44 - Castelfranco 153. 45- 47: S. gibbosa. 45 - JBL-04956; 46 - JBL-07529; 47 -<br />

JBL-07529. 48 - S. confusa (JBL-09484). 49 - S. grandiflora (JBL-09988). 50, 51 - S. ocul<strong>at</strong>a (Rojas 134; Rojas<br />

135). 52, 53 - S. x fowliana (JBL-1303l; JBL-1303l). 54 - S. costaricensis x cirrh<strong>at</strong>a (Rojas 113). 55- 65:<br />

Stanhopea spp. 55 - JBL-05654; 56 - FP. 2960; 57 - JBL-09988; 58 - FP. 2960; 59 - AK 557; 60 - FP 6279;<br />

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<br />

TABLE 2. Continued.<br />

dyana (3), S. panamensis, S. picta (5), Warczewiczella<br />

discolor (2), W. ionoleuca (2)<br />

Vandeae<br />

Aeridinae: Aerides falc<strong>at</strong>a, A . multiflora, Asco centrum<br />

curvifolium, Gastrochilus calceolaris, G.<br />

formosanum, Haraella retrocalla, Hygrochilus<br />

parishii, Phalaenopsis amabi[js (2), P. lueddemanniana,<br />

P. schilleriana, P. violacea, Sedirea<br />

japonica, Trichoglottis sp., Vanda coerulea, V.<br />

lamell<strong>at</strong>a<br />

Angraecinae: Angraecum ebumeum, A. eichlerianum<br />

Aerangiclinae: Eurychone rothschildiana<br />

Unplaced subtribes within Epidendroideae<br />

Dendrobiinae: Bulbophyllum lobbii, Dendrobium<br />

antennu<strong>at</strong>um, Epigeneium cymbidioides<br />

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