Rev. Biol. Trop. 46(4): 961-1031, 1998 www.ucr.ac.cr www.ots.ac.cr www.ots.duke.edu

Orchid florula of Parque Nacional Manuel Antonio, Quepos, Costa Rica

Franco Pupulin

Research Associate, Marie Selby Botanical Gardens. Research Associate, Jardín Botánico Lankester, Universidad de Costa Rica. Pennanent address: Vía della Liberta 17, I-21051 Arcisate (VA), Italy

Received 4-VI-1998. Corrected 31-VIII-1998. Accepted 10-IX-1998.

Abstract: An orchid florula of Parque Nacional Manuel Antonio, Costa Rica, is provided on the basis of collections made in 91 p1ots within the park and field observations carried out in the neighbouring areas; 39 species belonging to 25 genera are recorded. Orchid species are described and a key to the species for field identification, based on vegetative features, is provided. Analytica1 drawings are provided for 35 of the taxa considered. Ecological notes and estímate of frequency for each species within the Park are given. A new orchid species, Lockhartia pandurata, is described and illustrated.

Key words: Orchidaceae, Taxonomy, Frequency, Tropical wet forest, Costa Rica, Parque Nacional Manuel Antonio.

The Parque Nacional Manuel Antonio in the province of Puntarenas, about 175 km (PNMA) constitues a relict of the Pacific, coastal southwest of San José (9° 21' - 9° 25' N and wet forest in Costa Rica and a unique sample for 84° 06' - 84° 10' W) (Bolaños et al. 1983, the country of transitional life zone between Campos et al. 1983). The national park is tropical wet and tropical very wet forests (Tosi characterized by low hills and irregular 1969, Holdridge 1982). In spite of its reduced topography, with altitudinal range varying size, PNMA plays a significant role as an from sea level to about 150 m. Rainfall educational center, having one of the highest average per year ís 3,287 mm. Rainy months rates of visits of the whole Costa Rican park are from June through November, and the dry system both by local and foreign visitors. A first season extends from December to May (Rojas attempt to classify floristic and faunistic et al. 1983). Average temperature is 26.2 "C, components of PNMA was done in 1983 with 21 OC and 32 OC minumum and maximum (Bolaños et al.), but more specific studies are respectively. According to Rojas et al. (1983), needed in order to understand the actual richness the vegetation of PNMA may be subdivided in of the protected area. The hope of the present the following units: primary forest (105 ha), paper, intended to clarify the diversity of the disturbed primary forest (130 ha), secondary orchid farniliy at PNMA, is to represent a frrst forest (122 ha), young secondary forest (228 step for a systematic revision of the Park flora. ha), secondary forest with fruit trees (6 ha), mangrove (18 ha), pasture (47 ha), grassy MATERIALS AND METHODS lagoon (14 ha), and rock islands (13 ha) (Fig. 1). A slightly modified vegetation map of the Study area: The PNMA covers an area of central area of PNMA was done by Wong 683 ha along Costa Rican central Pacific coast Reyes (1990). Por the purpose of the present 962 REVISTA DE BIOLOGIA TRpPICAL

; /'--...... , -".. ··~·., . ~ N .. ..,.. .. ._. .. +10° 84° 374+ r-~ .. t..., r-' \ .J \

()

370+ 447 + 449 o 0.5 Km

- Primary forest 00 Pastores - Disturbad prirnary forest ~ Mangrove h>W;l Secundary torest D Beach ~ Young secundary torest ~ Rocks

Fig. l. Vegetation units of the park.

' . ... ' '• PUPUUN: Orchid Flon!la of Parque Nacional Manuel Antonio 963 study sjx units t.vere s,ampled. Beach Rican major herbaria were revised for orchid veget3.tion was incíuded in secondary mature material from the studied and neighbouring forest. Pastures were sampled through relictual areas. Nomenclature follows Dressler (1993). trees, that in sorne cases host a consistent Collections are deposited at the Herbarium of epiphytic flora due to thcir age. the Escuela de Biología, Universidad de Costa A total of 346 species of vascular Rica (USJ), whereas duplicates are kept at the cryptogams and angiospenns has been reported Herbario Nacional de Costa Rica (CR) and the for PNMA (Boza 1986). A preliminar list of Marie Selby Botanical Gardens (SEL). Orchidaceae, comprising 33 species in 23 genera, was done in 1983 (Bolaños et al.). However, apparentedly no collections were made at that RESULTS time, and no orchid material from this area is conserved eíther at the Herbario Nacional de The Orchidaceae constitute the largest Costa Rica (CR), the Herbarium of the Escuela family at PNMA, followed by Graminae and de Biología, Universidad de Costa Rica (USJ), or Leguminosae with about 25 and 23 species at the Instituto de Biodiversidad in Heredia respectively (Bolaños et al. 1983). 39 orchid (INB). Only two specimens determined as species, pertaining to 25 genera have been Lockhm1ia micrantha from Manuel Antonio and recorded. The genera richest in species are Punta Quepos, both collected outside the Park, Epidendrum (6 species) and Maiíllaria (4 are kept at USJ and CR. species), reflecting the very high number of Methods: The present list of orchid species within these genera all over the species is based upon collections gathered by country (Mora-Retana & Garcia 1992, the author during the months of June to August Dressler 1993). For each of the 39 orchid 1995. Collections were made in 91 plots ( 100 species frequency at PNMA has been assessed m2 each) of tropical wet, transition to tropical and classified. Species occurring in 25 or very wet forest at PN1'.1A, at intervals of about more 100 m2 plots are considered "very lO m between O and 150 m a.s.L (Fig. 2). common". Species present in 14 to 24 plots Species that were not collected at PNMA but are classified as "common". Taxa recorded in found by the author in natural populations in 6 to 13 plots are considered "ftequent". Those lowlands and hills around the Park are a1so found in 2 to 5 plots are regarded as included and marked in the key with an "occasíonal". Species observed in 1 plot or asterisk (*). The distance from PNMA is less only recorded from outside the Park area, are than 5 kilometers, and many of the taxa found considered to be "rare". Distribution of total at the Park form natural populations in these number of species over frequency classes is areas. Distribution of the plots resulted as shown in Fig. 3. "Very common" species follows: 17 in primary forest (18%), 14 in include Epidendrum amparoanum (44 ), disturbed primary forest (16%), 30 in Catasetum maculatum. (35), Scaphyglottis secondary forest (33% ), of which 8 plots in stellata (31), and Brassavola nodosa (26). beach vegetation, 22 in young secondary forest "Common" species include Dimerandra (24% ), 1 in secondary forest with fruit trees emarginata (21), Epidendrum congestum (21),. (1% ), and 7 isolated trees in pasture (8% ). Caularthron bilamellatum (15), and Within each plot orchid species were visually Lockhartia pandurata (16). "Frequent" determined, and two to three samples for each species ínclude Epidendrum stamfordianum apparentedly different taxon were collected, (6), Maxillaria neglecta (8), M. ponerantha together with additional data about (7), Nidema ottonis (6), and Pleurothallis vegetational aspects. Sexual material was corniculata. (9). "Occasíonal" and "rare" identified, and sterile material was cultivated species account for nearly 60% of total and flowered in Italy for identification. Costa species within the study area. 964 REVISTA DE BIOLOGIA TROPICAL

o o o \

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Fig. 2. Ubication of the plots within the study area. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 965

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40%- ---

30%- ---

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¡ 5 ~ o = ·¡;; .,O"' i i u o o "'u ~ u u o t:' ~ Fig. 3. Distribution of total number of species over frequency classes.

Distribution and number of orchid taxa distribution in young secondary forest reflects within the different types of vegetations preferences of sorne orchid species for disturbed strongly confirms the importance of habitats. lonopsis sa,tyrioides (Sw.) Rchb.f. and conservation of pristine habi~ats. Orchid Leochilus . scriptus (Scheidweiler) Rchb.f., as diversity is highest in primary forest, well as members of the genera Notylia and decreasing towards young secondary forest Trizeuxis, are known to ·be early colonizers of and beach vegetation (Fig. 4). Among the young canopies, and strong ecological evidences thirteen plots where no orchid species were were offered to explain site preferences for the observed, ten plots pertained to secondary and oblígate twig epiphytes (Chase 1987). young secopdary vegetation units. On the Notwithstanding, plots in young secondary contrary, all the five plots where the highest forest were relatively poor in orchid taxa, and number of orchid species was found ( 10 to 8 more than twenty species are found elsewhere in orchid species per plot) were located m the Park except in the areas covered with young primary and disturbed primary forest. secondary vegetation (Fig. S). The puzzle of different vegetation units within the Park also reflects on its orchid flora. DISCUSSION Species distribution within plots established in different vegetation units supports the view of The distribution of six species proved to be strongly ecological preferences for many of restricted to primary undisturbed forest, and this the orchid populations found at PNMA. Three figure raises to eight when also the plots of main groups may be identified. A first group is disturbed primary forest are included. However, represented by orchid species that form large it is interesting to note how a number of orchid populations in all types of vegetation, with a species are also restricted to secondary forest general preference for noderately to (six species) or young secondary forest (four completely exposed epiphytic situations. This species) (Fig. 5). It is likely the restriction oftaxa group includes common orchids like !1 !1 ~ (!<:¡ (!<:¡ "' V. ~ ;>::j o "' ;:¡ :J."' 1 2: n t0 o. ¡..,....,t. ,...... ~ ¡..-¡. ¡.....~. N o· o o o N .¡:.. o- 00 o. O N .¡:.. o- 00 O N .¡:.. o-, 00 O :::> 1 1- 1 =2' ...,o 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ~."' ¡;; 1 1 1 1 1 1 1 q >< 1 1 1 1 1 '1 1 1 .., 1 1 1 1 1 1 1 1 1 "' 1 1 1 1 1 1 1 ~. fr Species : 1 1 1 1 1 1 1 1 o restricted to : 1 1 1 1 1 1 1 1 'O "' 1 1 1 1 1 1 1 .., Primary "' 1 1 1 1 1 " undisturbed 1 1 <: S."'" 1 1 1 1 1 ro forest 1 1 1 1 1 1 (!<:¡ o 1 (j) primary forest : 1 1 1 1 1 "'o 1 1 1 1 g <: 1 1 1 1 1 1 e;· .., 1 1 1 1 1 1 1 Si o." 1 1 1 1 1 1 1 1 "' Species 1 1 1 1 1 1 1 1 < 1 1 "' ~ 1 1 1 1 E.· Disturbed Cñ ;;: restricted to : 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 primary forest ~ g primary 1 1 <: forest : á\ ()<:¡ 1;1:1 "1'1> g o· o~ Species 1 "' 1 1 1 1 i 1 Cl 1 1 1 1 1 1 1 Secondary ~·"' restricted : 1 1 1 ;; ;o ! ! 1 ! forest to secondary ~ forest o '"tJ Species ~ restricted to 1 1 1 1 1 1 1 1 1 1 1 1 1 Young young 1 1 1 1 1 1 1 1 1 1 1 secondary forest secondary 1 1 1 1 1 forest - Species distributed 1 1 elsewhere 1 1 Beach vegetation exceptin young secondary forest 1 PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 967

Brassavola, Catasetum, Caularthron, 2 (l).Leaves cylíndric (subterete to terete) ...... 3 Dimerandra, sorne Epidendrum and 2. Leaves flattened ...... :...... , ..... ;...... 5

Maxillaria species, Nidema, and 3 (2) Pseudobulbs cylíndric, more than 3 cm long ...... Scaphyglottis, all of them widespread along ...... ,...... ,..... Brassavola nodosa Pacific lowlands from southem Mexico to 3. Pseudoblbs rounded, very reduced, less than 1 cm Colombia. A second group is composed by long ...... 4 orchid species restricted to the remnants of 4 (3).Plants usually less than 10 cm long ...... primary or secondary mature forest, like ...... lonopsis satyrioides * Aspasia, Dichaea, Epidendrum 4. Plants usually more than 50 cm tal! (to 1 m) ...... stamfordianum, Lockhartia, Maxillaria ...... ;...... Oncidium dsceiulens ponerantha, Oncidium spp., Pleurothallis, Sobralia, Stenorrhynchos, Trigonidium, and 5 (2).Leaves laterally flattened ...... 6 5. Leaves dorsoventrally flattened ...... 7 Vanilla. Though sorne of the taxa within this group are of wide distribution in Mesoamerica, 6 (5).Sterns elongated, plants braídlíke ...... it also includes species supposedly endemic of ...... Lockhartia pandurata Costa Rican central Pacific lowland forests. 6. Sterns short; plants fan!íke ...... Trizeuxis fa/cata * Most of these species are shade tolerant 7 (5).Plants wíth pseudobulbs ...... 8 epiphytes, mainly established on trunks. and 7. Plant without pseudobulbs ...... , ...... 27 large axes of the phorophytes, where thick layers of bryophytes prevent complete 8 (7).Pseudobulbs arising frorn top of old pseudobulbs, dessiccation of the · substrate and the roots fortníng chaíns ...... 9 during the dry season. Along the borders of 8. Pseudobulbs arising frorn base of old pseudobulbs, PNMA 1 found representatives of a third group not forming chaíns ...... 1O of orchids, mainly composed by specialized 9 (8).Pseudobulbs pyriforrn, wider near the base ...... twig epiphytes, including Bulbophyllum, ...... Scaphyglottis stellata Campylocentrum, /onopsis, Leochilus, 9. Pseudobu1bs cylíndric, not wider near the base ...... Notylia, Polystachya and Trizeuxis. All of ...... Scaphyglottis prolifera * these orchids are rarely or only occasionally 10 (9).Leaves scattered along pseudóbulb ...... l1 found in the park. Seed morphology (Chase 10. Leaves (1-2) terminal on pseudobulb ...... 13 1997, Chase & Pippen 1988) and stress tolerance of juvenile twig epiphytes enabled 11 (10) V. Inflorescence Iateral ...... Catasetum maculatum them to establish and survive in the harsh 11. Inflorescence terminal ...... 12 conditions of young canopies and isolated ·12 (11).Pseudobulbs solíd ...... ;...... :...... trees in severely disturbed and .anthropic ...... Polystachya foliosa* habitats. Jeopardized vegetation with scattered 12. Pseudobulbs hollow ...... Caularthron bilamellatum phorophytes (e.g. orchard trees, trees in pastures, palm trunks, bushes along roadsides) 13 (10). Pseudobulbs rnonophyllous ...... 14 13. Pseudobulbs 2-leaved at apex ...... 18 support this way the dispersa! of specialized members of the Orchidaceae that are 14 (13).·Plants cespitose (rhizorne very short) ...... 15 progressively integrating the orchid stock of 14. Plants wíth pseudobulbs scattered on a long rhizorne.l7 the original forests. 15 (14).1nflorescence penduloús ... c...... Notylia pittieri 15. Inflorescence erect ...... l6

KEY TO THE SPECIES 16 (15).1nflorescence lateral ...... Leochilus labiatus 16. Inflorescence termina! ...... Nidema ottonis

l. Plant terrestrial ...... Stenorrhynchus lanceolatum * 17 (14). Inflorescence erect...... Maxillaria oreocharis l. Plant epiphytíc ...... 2 17. Inflorescence pendulous ...... Maxillaria neglecta 968 REVISTA DE BIOLOGIA TROPICAL

18 (13). Pseudobulbs atless threetimes longerthan wide ...... 19 ...... Epidendrum congestum 18. Pseudobulbs not more than two times longer than 33. Stem more than 15 cm long ...... 34 wide ...... 21 34 (33). Stem with 4-6leaves only at apex ...... 19 (18). Pseudobulbs of a single intemode ...... Epidendrum nocturnum ...... Encyclia abbreviata 34. Stem covered by leaves for al! its length ...... 35 19. Pseudobulbs of more than one intemode ...... 20 35 (34). Base of the leaves completely envolving the stem 20 (19) Pseudobulbs less than 1O cm long; inflorescence ...... Epidendrum amparoanum terrninai ...... Scaphyglottis micrantha 35. Base ofthe leaves only partially envolving the stem. 36 20 . Pseudobulbs more than 20 cm long; inflorescence from the rhizome at the base of pseudobulb ...... 36 (35). Leaves elliptíc, obtuse, fleshy ...... Epidendrum stamfordianum ...... Epidendrum sculptum 36. Lea ves lanceolate, acute, not fleshy ...... 21 (18). Basalleaves (cataphylls) present...... 22 ...... Epidendrum isomerum * 21. Basalleaves ( cataphy lis) absent ...... 26 37 (31). Inflorescence 1-flowered .. Dichaea panamensis * 22 ( 19). Plant pendent ...... Maxillaria crassifolia 37. lnflorescence many-flowered ...... 38 22. Plant erect ...... 23 38 (37). Stem less than 20 cm; lea ves about 4 cm long ..... 23 (22).Plant small, about 5 cm tal! ...... Campylocentrum micranthum ...... Maxillaria ponerantha 38. Stem more than 30 cm long (50 cm): 23. Plant more than 15 cm tall ...... 24 leaves about 8-10 cm long ...... Campylocentrum panamense * 24 (23). Pseudobulbs club shaped.Aspasia epidendroides 24. Pseudobulbs not club shaped ...... 25

25 (24)Pseudobulbs widely elliptic, very flat...... LIST OF SPECIES ...... Oncidium stenobulbon * 25. Pseudobulbs o vate, sulcate, not flat ...... l. ASPASIA Lind 1 • ...... Oncidium polycladium *

26 (21).lnflorescence erect...... Trigonidium egertonianum Cespitose, epiphytic herbs with 1-2 leaved 26. Inflorescence pendent ...... Bulbophy/lum oerstedii * pseudobulbous stems. Leaves subcoriaceous at apex of the pseudobulb and along the stem. 27 (7) Planta vine ...... Vanilla pompona * lnflorescence a lateral many-flowered, often 27. Plant not a vine ...... 28 successive, spicate raceme. Flowers rather 28 (27). Single leaf on each stem ...... 29 large and showy, greenish-brown stripped with 28. Leaves severa! on each stem ...... :.30 brown. Sepals and petals subequal, free: lip with a claw entirely adnate to the column and a 29 (28). Leaf coinlike, rounded, flat on the bark ...... broad lamina with severa! keels. Column erect, ...... Pleurothallis lewisae 29. Leaflanceolate, not flat on the bark ...... nearly terete, footless, wingless. Pollinia 2...... Pleurothallis corniculata A genus of about 6 species native of Central and South America. 30 (29). Leaves distinctly pleated ...... Sobralia decora 30. Lea ves not pleated ...... 31 l. Aspasia epidendroides Lindl., Hook.

31 (30). Inflorescence terminal ...... 32 Jour. Bot. 1: 6. 1834. Aspasia fragrans Kl., 31. Inflorescence lateral ...... 37 Ind. Sem. Hort. Bot. Berol.l2. 1852. Odontoglossum aspasia Rchb.f., Walp. Ann. 6: 32 (31 ). Plant with thickened stem ...... 851. 1861. (Fig. 6) ...... Dimerandra emarginata Plant epiphytic, medium sized, to 35 cm 32. Plant with slender stem ...... 33 tall, with terete rhizome and rather spaced 33 (32).Stem less than 5 cm long, completely concealed by pseudobulbs. Roots fleshy, with green the base of the lea ves ...... vegetative apex. Pseudobulbs diphyllous, PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 969

F

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10 cm

3 cm

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Fig. 6. Aspasia epidendroides Lindl. A- Habit. B - Flower. C - Column and lip, lateral view (sepals and petals removed). D - Dissected perianth. E - Column, ventral view. F. Pollinarium and operculum. Illustration voucher: F. Pupulin 311. 970 REVISTA DE BIOLOGIA TROPICAL stlpttate, oblong-elliptic to linear-elliptic, conduplicate to terete, linear leaves. laterally compressed, 5-12 cm long, 3-6 cm wide, Inflorescence a terminal or lateral 1- to many­ the cylindric lower portions covered with several flowered raceme. Flowers large and imbricating bracts, the uppermost 2 or 3 conspicuous, white. Sepals and petals foliaceous. Leaves elliptic-lanceolate, subequal, free; lip with a tubular claw enfoding subcoriaceous, acute, up to 24 cm long, 2.5-5 cm the column and a broad lamina. Column very wide. Inflorescences 1-2 simple, erect, few­ short, footless, winged. Pollinia 8. flowered racemes, to 18 cm long, produced from A neotropical genus of about 17 species. the axils of the uppermost foliaceous bracts; bracts ovate-lanceolate, about 1 cm long. 1. Brassavola nodosa (L.) Lindl., Gen. et Flowers showy, scented, the sepals green with Sp. Orch. Pl. 114. 1831. Epidendrum nodosum broad transverse brown bands, the petals pale L., Sp. Pl. 953, 1753. Brassavola venosa lavender to grenish with transverse bars, the lip Lindl., Bot. Reg. 26: Mise. 20, 1840. white with conspicuous la vender markings and a Brassavola scaposa Schltr., Orchis 13; 77, yellow callus, the column tinged with lavender. 1919. (Fig. 7). Sepals elliptic to elliptic-obovate, acute, Plant epiphytic, medium sized, up to 50 obscurely apiculate, to 2.8 cm long, 0.9 cm wide, cm tall, with short, terete rhizome. Roots the lateral sepals often reflexed. Petrus elliptic­ fleshy, with green to brownish vegetative apex. obovate, concave, obscurely keeled at the dorsal Pseudobulbs terete to davate, slender, 2.5 to apex, to 2.1 cm long, 0.9 cm wide. Lip clawed, 8.5 cm long, about 0.6 cm wide, covered by 2- to 3 cm long, 2.4 cm between the lateral lobes; 3 scarious, tubular. sheaths, monophyllous. the claw adnate to the lower portian of the Leaf conduplicate, very fleshy-coriaceous, column, about 8 mm long, the subquadrate, linear, acuminate to slightly mucronate, sulcate obscurely 3-lobed lamina abruptly deflexed, the above, up to 20 cm long, 0.9-1.2 cm wide. rnidlobe emargínate with undulate margins, the Inflorescence terminal, to 13 cm long, 1- to disk with 2 parallel extending from the base to many-flowered (mainly 2-fowered); peduncle the insertion of the rnidlobe. Column semi terete, terete; bracts triangular-lanceolate, acute, 0.7-1 basally adnate to the lip, to 2.3 cm long. Pollinia cm long. Flowers large and showy, with pale 2, obpyrifonn, on a short ligulate stipe; green sepals and petals, and white lip spotted visciudium elliptic. Anther cap herniglobose, by pale purple at the base. Sepals subequal, keeled. linear-lanceolate, acute, to 9 cm long, 0.6 cm Distribution: From Guatemala to Panama. wide. Petals linear, attenuate, to 8.1 cm long, Widespread but not common at low elevations 0.4 cm wide. Lip with a tubular claw, abruptly in Central America. expanding into a ovate-cordate lamina, Parque Nacional Manuel Antonio: apiculate at apex, to 7.8-8 cm long, 4.3 cm Epiphytic on lower branches and trunks of wide at mid-point of the lamina; claw about 3 trees mainly in primary or mature secondary cm long, 1.2 cm wide, with erose-dentate forest. Plants are usually found in shaded margins. Column short, with a pair of conditions. Occasional at PNMA. triangular-subfalcate, acute wings, 3-lobed at Phenology: flowering mainly occurs in apex, to 7 mm long. Pollinia 8, in two August-September. It rnay also flower from subequal groups of 4, on two powdery, bilobed February to April. caudicles. Anther cap hemiglobose, 8-celled. Distribution: From Mexico to Panama and northern South America. Common at lower 2. BRASSAVOLA R. Br. elevatíons on Pacific coasts in Central America. Cespitose, epiphytic or lithophytic herbs Parque Nacional Manuel Antonio: with terete, monophyllous pseudobulbs and Epiphytíc on Hippomane mancinella PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 971

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Fig. 7. Brassavola nodosa (L.) Lindl. A- Habit. B - Flower. C - Column and lip, lateral view (the lip longitudinally sectioned, sepals and petals removed). D - Dissected perianth. E - Column, ventral view. F. Operculum and pollinaria. Illustration voucher: F. Pupulin 331. 972 REVISTA DE BIOLOGIA TROPICAL

(manzanillo), Tabebuia rosea (roble de many-flowered, the peduncle slender, sabana), Cocos nucifera and, less frequently, becoming fleshy at the rhachis, 12-25 cm long; on Terminalia catappa (almendro de playa) bracts broadly triangular, acute, 0.3 mm long. trees just to the line of seashore. Also epiphytic Flowers small and inconspicuous, subsessile, in mangrove vegetation on Rhizophora sp. born in shallow depressions on the rhachis, Plants of B. nodosa are usually growing in paJe green to yellowish green finely spotted exposed to medium-shaded conditions. White with purple; lip purple at apex. Dorsal sepal color of the flowers without any nectar guide free, concave, ovate, acute, 6 mm long, 3.5 mm and smell at night are characters generally wide. Lateral sepals ovate-acuminate, associated to pollination by sphingid moths. sometimes slightly connate atthe base, adnate However, the actual pollinator of the species is to the foot of the column, 7 mm long, 3 mm still unknown. Very common at PNMA. wide. Petals oblong elliptic, obtuse to Phenology: flowering mainly occurs in subacute, 2.5 mm long, 1 mm wide. Lip entire, July-August. ligulate, obtuse, very fleshy toward the apex, , geniculate in natural posistion, contracted at the base and articulate to the foot of the 3. BULBOPHYLLUM Thouars column, 2.5 mm long, 1 mm wide. Column subterete,3-lobed at apex, provided with a Epiphytic herbs, usually with 1-2 leaved distinct foot at the base, about 1.8 mm long. pseudobulbs. Leaves subcoriaceous at the apex Pollinia 4, in two cohalescent pairs. Anther of pseudobulbs. Inflorescence lateral from the cap operculate, triangular, 1-celled. base of the pseudobulb, a spike, raceme and Distribution: Widespread but not common rarely a single flower, sometímes thickened or in lowlands in Central and South America, this widely flattened. Flowers often with a mimicry species has been widely known as B. and colours intended for flies pollination. pachyrhachis, possibly an endemic ofthe West Dorsal sepa! free, the laterals often connate Indies. and adnate to the column foot; petals usually Parque Nacional Manuel Antonio: A smaller than sepals, variously ornate; lip thin single plant of this species was found on fruit or rigid, usually fleshy, articulated to the tree not far from the eastern border of PNMA. column foot. Column erect, auriculate, The plant was growing on a large branch forming a foot at the base. Pollinia 4. partially covered by mosses in medium shade. A pantropical genus, mainly Asiatic, of It should be considered rare at PNMA. about 1,000 species. Phenology: flowering occurs in December and January. l. Bulbophyllum oerstedii (Rchb.f.) Hemsley, Biol. Centr.-amer. 3: 213. 1883. Bulbophyllaria oerstedii Rchb.f., Bonpl. 3: 4. CAMPYLOCENTRUM Benth. 223. 1855. Bulbophyllum pachyrhachis Auct., non Grisebach. (Fig. 8). Epiphytic monopodial herbs with either Plant epiphytic, medium sized, up to 20 distichous, leafy stems or leafless condensed cm tall. Roots filiform, glabrous. Pseudobulbs stems. Leaves oblong or ligulate, coriaceous, subconical, strongly 4-angulate, slender, 2.5 to sometimes absent. Inflorescence a short 4 cm long, about 2.5 cm wide, widely spaced raceme, usually densely flowered, produced on a creeping rhizome, covered at the base by from the axils of the leaves or from the short 2-3 scarious, tubular sheaths, diphyllous. stem at the center of the root cluster. Flowers Leaves coriaceous, linear-lanceolate, acute to often distichously arranged on the rhachis. acuminate, up to 20 cm long, 1-2.5 cm wide. Sepals subequal, free or partially connate; Inflorescence lateral, arching to pendent, petals subequal to the sepals; lip sessile, 1-3 PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 973

3mm

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smm 10 cm.

Fig. 8. Bulhophyllum oerstedii (A. Rich. & Gál.) Hamer & Garay. A- Habit B - Flowers. C- Column and lip, laterál view. D- Dissected perianth. E- Column, ventrál view.F- Apex of colulnn, ventrál view (anther removed). G. Pollinia and operculum. lllustration voucher: F. Pupulin 453. 974 REVISTA DE BIOLOGIA TROPICAL lobed, forming an elongate or saccate spur at situations with their adventitious roots loosely the base. Column wingless, without a foot. anchored to the host twigs. Rare at PNMA. Pollinia 2. Phenology: flowering mainly occurs in A neotropical geous of about 50 species. July-August.

l. Campylocentrum níicranthum (Lindl.) 2. · CampyliJcentru~ panamense Ames, Rolfe, Orch. Rev. 11: 245. 1903. Angraecum Or~hidaceae 7; 88:,1922. (Fig. 10). micranthum Lindl., Bot. Reg. 21: t. 1772. Plant epiphy~ic; n:ú:mopodial, pendent, 1836. Angraecum brevifolium Lindl., Bot. with terete, leafy st~m up to 50 cm long. Roots Reg. n.s. 3: sub t. 68. 1840 .. Angraecum fleshy, altemating to the leaves. Leaves elliptic lansbergii Rchb.f., Nedéri. Kruidk. Arch. 4: to linear-elliptic, distichous, emarginate, 316. 1859. Aefanthus ·miÚanihus · Rchb.f., unequally 2-lobed a.t apex, articulate with the Walp. Ann. 6: 901. 1864. Aeranthus lansbergii sheath envolvig the stem, datk green in color, Rchb.f., Walp. Ann. 6: 901. 1864. to about 8.5 cm long, 2.5 mm wide. Campylocentrum peniculus Schltr., Fedde Inflorescence a raceme shorter than the Rep. Spec. Nov. Beih. 17: 91. 1922. (Fig. 9). leaves, produced along the stem, with the Plant epiphytic, monopodial, small, with flowers arranged distichously on the rhachls, terete stem to 20 cm long, leafy above. Roots about 5 cm long. Flowers small, greenish­ fleshy, alternating to the leaves. Leaves white with the spur tinged by ·. yellow, the eliptic to suborbicuar, distichous, coriaceous, segments not well spreading. Dorsal sepal obtuse, unequally 2~lobed at apex, articulate linear-elliptic, obtuse, concave towards the with the tubular sheath envolving the stem, base, 5 mm long, 1.5 mm wide. Lateral sepals pale green i~ color, to. about 5 cm lorig, 2-2.5 linear-spathulate, wider near the base, acute, cm wide. Infloréscence araceme produced with a small apicule, com;ave, about 6 mm from the stem below the leaves, usually long, 1.3 mm wide. Petals similar to the dorsal shorter than the leaves; with the flowers sepal, to 4.5 mm long, 1.5 mm wide. Lip arranged distichously on the rhachis. Floers obscurely 3-lobed, the laterallobes triangular, small, white with the sur tinged by yellow, the small, rounded at apex, clasping the column, segments not well spreading. Sepals the midlobe lanceolate, acute, concave, subequal, lanceolate, acute, concave towards sparsely covered with hairs along the midrib, the base, to 5 mm long, 1.5 mm wide. Petals to 5 mm long, 2.2 mm wide between the lateral similar to the sepals, to 4.5 mm long, 1.3 mm· lobes, extending at the base into a cylindric, wide. Lip 3/lobea; the lateral lobes broadly . slightly curved spur, more or less continuous triangular, rounded at apex, clasping the with the lip, abou.t 5.5 mm long. Column midlobe triangular/ la~ceolate, acute,' short, o.s mm long. Pollinia 2 on short ligulate concave, sparsely covered with short airs, toS · stipes. Anther cap subquadrate, 2-celled. mm long, 2 mm wide between. the lateral Distribution: P¡;obably endemic in Costa lobes, extending at the ·base into a Rica and Panama, but plants of thls species cylindric/clavate, down/tumed spur about 4 may have been confused elsewhere with the mm long. Column short, about 1 .inm long .. widespread C. micranthum. Pollinia 2 on short ligualte stipes. Anther Parque Nacional Manuel Antonio: So far cap subquadrate/cucullate, 2/celled. the species has not yet been collected within Distribution: Widespread in lowlands the Park area. A population was found in a throughout the neotropics, the species may be riparian secondary forest on hills sorne 10 km locally abundant. West of PNMA. The plants grows on branches Parque Nacional Manuel Antonio: Epiphytic covered by mosses directly hanging over a on short trees in yoÚng secondary forest, where little stream in shade. It should be considered the plants are usually observed in exposed rare at PNMA. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 975

3 mm.

1 mm

5mm

Fig. 9. Cqmpylocentrum micranthum (Lindl.) Rolfe. A- Habit. B - Flower. C-Lip with spur, lateral view (sepals and petals removed). D - Dissected perianth. E - Operculum. llustration voucher: F. Pupulin 332. 976 REVISTA DE BIOLOGIA TROPICAL

10 cm

5mm 1 cm

Fig. 10. Campylocentr~t~r~ panamense Ames. A- Habit. B - Flower. C - Lip with spur, lateral víew (sepals and petals removed). D- Dissecled perianth. E- Operculum and pollinarium. Illustratíon voucher: F. Pupulin 305. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 977

Phenology: flowering occurs in Staminate flowers (male) yellowish-green, September and October. suffused and spotted by purple. Dorsal sepal membranaceous, oblong-elliptic to lanceolate, acuminte, concave, to 4.2 cm long, 1.6 cm 5. CATASETUM L.C. Rich. ex Kunth wide. Lateral sepals membranaceous, obliquely lanceolate, acuminate, to 4.4 cm Epiphytic or rarely terrestrial herbs with long, 1.9 cm wide. Petals membranaceous, ovoid or fusiforrn, many-leaved pseudobulbs. elliptic-lanceolate, acuminate, mucronate at Leaves usually large and plicate, deciduous. apex, 4 cm long, 1.8 cm wide. Lip fleshy, Inflorescence lateral, from near the base of the rigid, saccate, obconic, 3 cm long, 2.5 cm pseudobulb, a many-flowered raceme. Flowers wide, margins of the basal portian of the usually large and conspicuous, unisexual, orífice ciliate, apex of the orífice slightly dimorphic, non-resupinate. Sepals and petals emarginate. Column stout, rostrate, concave, subequal, free, fleshy or membranaceous; lip 3.2 cm long, 1.2 cm wide, with 2 slender, thin or rigid, sessile, lateral compressed on elongate antennae projecting downward, one deeply concave. Column erect, thickened, with unciforrn, the other undulate, extending into or without a pair of antennae at the base in the calceolate lip. Pollinia 2, obovate, sulcate, male flowers; shorter and thicker, without on a obtriangular deflexed stipe; viscidium antennae in female flowers. Pollinia 4. peltate. Anther cap cucullate-rostrate, 2- A genus of about 100 species restricted to celled. Pistillate flowers (female) yellowish the tropical regions of America. green, spotted by purple at the base of petals. Sepals and petals reflexed, fleshy. Sepals l. Catasetum maculatum H.B.K., Syn. subequal, broadly ligular, apiculate at apex, to PI. Aequinoct. 1: 331, 1822. Catasetum 2.8 cm long, 2 cm wide. Petals elliptic­ oerstedii Rchb.f., Bonplandia 3: 218, 1855. lanceolate, acute, to 2.5 cm long, 1.9 cm wide. Catasetum rostratum Klinge, Acta Hort. Lip fleshy, rigid, saccate, 3.1 cm long, about 2 Petropol. 17: 134, t. 2, 1898. Catasetum cm wide, margins of the orífice smooth. brenesii Schltr., Beitr. Ork. Zentralam. 2: 136 Column fleshy, very stout, to 1.4 cm long, 1.2 et 225, 1923. (Fíg. 11). cm wide, with a short apicule on upper apex. Plant epiphytic, ascending, stout, with Distribution: From Mexico to Colombia. abbreviated rhizome, to 70 cm tall. Roots Rather common at lower elevations in Central fleshy, glabrous, with many slender, erect, America. secondary rootlets, forrning dense mats around Parque Nacional Manuel Antonio: the plant base. Pseudobulbs subfusiforrn­ Epiphytic on Tabebuia rosea (roble de sabana), conical, to 30 cm long, 5-8 cm wide, covered Hymenaea courbaril (guapinol), Cocos by 5-6 scarious, imbricating leaf-sheaths. nucifera and Terminalia catappa (almendro de Leaves 6 to 8, distichous, elliptic-lanceolate to playa) trees just to the line of seashore, oblong-elliptic, acute to acuminate, plicate, generally exposed to full sunlight. Pollinated strongly nerved, up to 45 cm long, 8-12 cm by the bee Eulaema cingulata (Allen 1952). wide, the blades deciduous, the persistent Common at PNMA. bracts enveloping the pseudobulbs armed at Phenology: flowering mainly occurs in apex with sharped spines. Inflorescence July-August. lateral, basal, erect to arching, racemose, 3-to 14-flowered, up to 40 cm long. Ovaries pedicellate, stout, arcuate, to 3 cm long. 6. CAULARTHRON Rafin. Flowers relatively large, non-resupinate, unisexual, dimorphic, staminate and pistillate Epiphytic herbs with fusiforrn, usually flowers produced on separate inflorescences. hollow pseudobulbs inhabited by ants, few-leaved 978 REVISTA DE BIOLOGIA TROPICAL

..____...... ::.3. cm

...... _...;.3_C':.l_ __J

'-----3 cm___ _, 3cm _1

Fig. 11. Catasetum maculatum Kunth. A - Habit. B - MaJe flower. C - Male column and lip, lateral view (the lip longitudinally sectioned, sepals and petals removed). D - Dissected male flower. E - MaJe column, ventral view. F - Pollinarium and operculum. G- Female flower. lllustration voucher: A-F, F. Pupulin 323; G, F. Pupulin 327. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 979 at apex. Leaves conduplicate, coriaceous. lobed, ovate-lanceolate, the obscure lateral lnflorescence terminal from the apex of the lobes widely triangular, the midlobe acute to pseudobulb, a many-flowered raceme. Flowers slightly apiculate, the entire lip 1.6 cm long, small or large and conspicuous. Sepals and 0.7 cm wide, provided at the base with two petals similar, free, spreading; lip 3-lobed, free small rounded lobules; disc with two large, from the column, with two horn-like, hollow lamellate, hollow, horn-like projections near projections near the base. Column short, flat, the middle. Column stout, concave, 9 mm without a foot, provided with lateral wings. long, with 2 wide, rounded wings at the Pollinia 4. middle. Pollinia 4, laterally compressed, each A neotropical genus of about 4 species. pair provided with a granular, viscid, claudicle. Anther cap cucullate. l. Caularthron bilamellatum (Rchb.f.) Distribution: From Guatemala to R.E. Schultes, Bot. Mus. leaf. 18: 92. 1958. Venezuela, Trinidad, and Ecuador. Rather Epidendrum bilamellatum · Rchb.f., Wa1p. common at lower elevations and in dry forests Ann. Bot. Syst. 6: 345. 1862. Epidendrum in Central and northern South America. bigibberosum Rchb.f., Walp. Ann. Bot. Syst. Parque Nacional Manuel Antonio: 6: 346. 1862. Diacrium bigibberosum Hemsl., Epiphytic on Tabebuiarosea (roble de sabana) Godm. & Salvin Biol. Centr.-Amer, Bot. 3: and Terminalia catappa (almendro de playa) 222. 1883. Diacrium bilamellatum Hemsl., trees, mainly close to the sea, in medium Godm. & Salvin BioJ. Centr.-Amer. Bot. 3: shaded situations or exposed to full sunlight. 222. 1883. Diacrium indivisum Broadway, Sometimes íhe flowers are self-pollinating Bull. Mise. Inform. Trinidad. 2: 79. 1895. before bud opening. Common at PNMA. Diacrium venezuelanum Schltr., Repert. Spec. Phenology: flowering mainly occurs in Nov. Regni Veg. Beih. 6: 41. 1919. Diacrium March-April. bilamellatum var. reichenbachianum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 17: 47. 7. DICHAEA Lindl. 1922, in textu. Diacrium bivalvatulum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 123. Epiphytic monopodial herbs, lacking 1923. (Fig. 12). pseudobulbs, pendent or rarely erect, with Plant epiphytic, ascending, stout, with foliaceous stem. Leaves conduplicate, short rhizome, to 50 cm tall. Roots fleshy, articulate or not, membranaceous or glabrous. Pseudobulbs cylindric to fusiform, coriaceous. Inflorescence lateral from the axils swollen, of severa! nodes, to 30 cm long, 2.5-4 of the leaves, 1-flowered on a short peduncle. cm wide, covered by many imbricating, pale Flowers small or medium sized. Sepals and green sheaths, turning withish grey with age, petals subequal, free or the laterals sepals 2-4 leaved at apex. Leaves elliptic to Iigulate­ sÜghtly connate, spreading; lip clawed, 3- elliptic, obtuse, fleshy, up to 18 cm long, 0.8- lobed or rarely entire. Column short, erect, 2.5 cm wide. Inflorescence terminal from the with a short foot, provided with a infastigmatic apex of the pseudobulb, a raceme erect, ligule. Pollinia 4, waxy. several-flowered, the flowers opening A neotropical genus of about 45 species, successively at the end of an elongate peduncle often restricted to the trunks and lower up to 40 cm long. Ovaries pedicellate, to 3 cm branches of the host trees. long. Flowers white, the apex of sepals suffused by rose-brown, often cleistogamous. l. Dichaea panamensis Lindl., Gen. et Sepals subequal, elliptic to elliptic-obovate, Spec. Orch. PI. 209. 1833. Epithecia acute, dorsally carinate, 1.5 cm lohg, 0.9 cm panamensis Schltr..• Orchis 9: 25. 1815. wíde. Petals broadly obovate, with a short Dichaeopsis panamensis (Lindl.) Schltr., Beih. claw, subacute, 1.5 cm long, 1 cm wide. Lip 3- Bot. Centralbl. 36(2): 519. 1918. (Fig. 13). 980 REVISTA DE BIOLOGIA TROPICAL

1 cm

3 cm

1 cm 2 cm

Fig. 12. Caulathron bilamellatum (Rchb.f.) R.E. Schult. A- Habit. B - Floer. C - Column and lip, lateral view (speals and petals removed). D- Dissected perianth. E- Column, ventral view. F. Pollinaria and operculum. lllustration voucher: F. Pupulin 355. PUPULIN: Orchid Aorula of Parque Nacional Manuel Antonio 981

E

2mm

5 mm 1 cm

Fig, 13. Dichaea panamensis Lindl. A- Habit. B - Aower. C - Colurnn and lip, lateral view (sepals and pe.als removed). D - Dissected perianth. E - Pollinariurn and operculurn. Illustration voucher: F. Pupulin 304. 982 REVISTA DE BIOLOGIA TROPICAL

Plant epiphytic, pendent, with subcoriaceous, scattered along the upper part monopodial, foliaceous stem, up to about 15 of the stems. Inflorescence terminal, a very cm tall. Roots fleshy, glabrous, wider than the short, few- successively flowered raceme. stem, about 2 mm in diameter. Leaves Flowers of medium size, showy, rose-purple arranged distichously on the stems, narrowly wíth white center. Sepals and petals subsimilar, linear-elliptic, acute and apiculate, spreading, the petals wider; lip simple with a subcoriaceous, 1.2-2 cm long, 0.3-0.4 cm short claw, provided with a callus formed by wide, the blade articulated to sheath envolvíng severa! obscure carines. Column short, the stem and ultimately deciduous. apically provided with two wings. Pollinia 4, Inflorescences lateral from axils of the lea ves, waxy. 1-flowered, on a short peduncle 1.5 cm long. A genus of about 8 species, often self­ Ovaries pedicellate, glabrous, about 2 mm pollinating, wídely distríbuted in drier forest long. Flowers greenish white spotted with throughout tropical America. violet. Dorsal sepal ovate-elliptic, acute, dorsally carinate, 6 mm long, 2.5 mm wide. l. Dimerandra emarginata (G.F. W. Lateral sepals ovate-lanceolate, slightly Meyer) Hoehne, Bol. Agric. (Sao Paulo) 34: falcate, acute, apiculate, 7.5 mm long, 3 mm 618. 1934. Oncidium emarginatum G.F.W. wide. Petals obliquely ovate, acute, 6 mm Meyer, Prim. Fl. Essequeb. 259. 1818. long, 4 mm wide. Lip 3-lobed, anchoriform to Epidendrum stenopetalum J.D. Hook., Bot. sagittate, with a fleshy claw, the lateral lobes Mag. 62: t. 3410. 1835. Epidendrum narrow, retrorse, acute, the midlobe broadly stenopetalum var. tenuicaule Rchb.f., Otía Bot. triangular, shortly acute, adaxially carinate hamb. 1: 12. 1878. Dimerandra stenopetala towards the apex, the entire lip 7 mm long, 6 (J.D. Hook.) Schltr., Repert. Spec. Nov. Regni mm wide between the lateral lobes. Column Veg. Beih. 17: 44. 1922. (Fig. 14). erect, with a short foot, provided at the base of Plant epiphytic, cespitose, with fleshy, the stigmatic cavity wíth a glabrous ligule, 2.5 foliaceous stem, up to about 60 cm tall. mm long. Pollinia 4, laterally compressed, in Pseudobulbs cylindric to fusiform from a two pairs superposed, on a triangular elongate narrow stalk, of severa! nades, bearing leaves stipe. Anther cap hemiglobose, 2-celled. in the upper portian. Roots fleshy, glabrous. Distribution: A very common epiphyte in Leaves arranged distichously on the stems, Central America. It is usually found in shade narrowly linear-elliptic, subcoriaceous, on trunks and branches of understory trees at unequally bilobed at apex, 4-9 cm long, 0.7- low elevations from Mexico to Panama. 1.1 cm wide, produced at the nades of the Parque Nacional Manuel Antonio: The pseudobulb. Inflorescences terminal, a short species has not yet been collected withín the raceme bearing 1- to few flowers produced Park. Plants were found at the base of a large successively about 1 cm long. Flowers very Ceíba sp., in primary forest remnants, on hills flat, often self-pollinating, rose purple with a sorne 1O km West of PNMA. So far, it should white blotch at the base of the lip. Sepals be considered rare at PNMA. subsimilar, elliptic-lanceolate, acute to Phenology: flowering occurs throughout acuminate, the lateral sepals somewhat the year. oblique, 15 mm long, 5 mm wide. Petals broad-obovate to subrhombic, acute, 17 mm long, 10 mm wide. Lip flabellate, with a fleshy 8. DIMERANDRA Schltr. claw partially adnate to the column, the lamina broadly obovate to suborbicular, truncate, Herbs epiphytic, cespitose, with slightly apiculate, the disc wíth a callus thickened, foliaceous stem. Leaves composed by obscure línes, the entire lip 16 conduplicate, linear to narrowly elliptic, mm long, 15 mm wide. Column short, without PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 983

F

5 mm

10 cm

2 cm

2 cm

Fig. 14. Dimerandra emarginata (G.F.W. Meyer) Hohene. A- Habit. B- Flower. C- Column and lip, lateral view (sepals and petals removed). D - Dissected perianth. E - Pollinaria and operculum. Illustration voucher: F. Pupulin 325. 984 REVISTA DE BIOLOGIA TROPICAL a foot, provided at the apex with two, large, Plant epiphytic, with elongate rhizome subquadrate, deflexed wings, about 5 mm and well spaced pseudobulbs, up to about 20 long. Pollinia 4, laterally compressed, in two cm tall. Pseudobulbs stalked, fusiform, pairs on short, bílobed caudicles. Anther cap slightly flattened, 4-5 cm long, to 0.9 cm herniglobose, 4-celled. wide, 2- to 3-leaved at apex. Roots filiform, Distribution: A common epiphyte in glabrous. Leaves narrowly linear-elliptic, soft, tropical America. lt is usually found at low minutely retuse, 4-15 cm long, 0.4-1 cm wide. elevations from Mexico to South America and Inflorescences terminal, a short raceme the West Indies. bearing 2-4 flowers, produced from a Parque Nacional Manuel Antonio: The inconspicuous spathe, about 1.5 cm long. species may be observed growing Flowers small, creamy white striped with epiphytically in any kind of vegetation at the violet on the lip. Sepals subsimilar, Park, the plants often forming big clumps of lanceolate, acute, the lateral sepals oblique, 13 many stems, usually pendent in mature mm long, 2.5 mm wide. Petals oblanceolate, specimén. Sometimes the flowers are self­ acute, 12 mm long, 2 mm wide. Lip with a pollinating soon after bud opening. Very short claw, adnate to basal half of the column, common at PNMA. flabellate, the lamina ovate, rounded to Phenology: flowering occurs from cordate basally, acute, the disc with a September through December. subquadrate thickening, 11 mm long, 7 mm wide. Column subterete, flattened, widened at the middle, 3-toothed at apex, the mid-tooth 9. ENCYCLIA Hook. shorter than the lateral ones, about 5 mm long. Pollinia 4, Jaterally compressed, in two pairs Epiphytic herbs with ovoid or elongated, on bilobed caudicles. Anther cap heteroblastic pseudobulbs, 1- to 3-foliate at hemiglobose, 4-celled. apex, the base envolved in papyraceous Distribution: From Mexico to Central sheaths. Leaves conduplicate, oblong to linear, America, usually in evergreen forest just to subcoriaceous to leathery. Inflorescence 1,000 meters elevations. terminal, sometimes produced from a small to Parque Nacional Manuel Antonio: The large spathaceous bract, a few- to many­ species has been found growing epiphytically successively flowered raceme or panicle. in secondary forest in shaded conditions. Rare Flowers from small to large and showy. Sepals at PNMA. subequal, free, spreading; the petals usually Phenology: flowering mostly occurs in similar to sepals or slightly narrower or May-June. broader; lip free from the column or adnate only at the base, simple or 3-lobed, provided with various kinds of calli. Column stout or 10. EPIDENDRUM L. slender, footless, often provided with apical wings. Pollinia 4, waxy. Small to large, ces pi tose or repent, erect to A neotropical genus of about 150 species, pendent, epiphytic or rarely terrestrial herbs to usually well represented in areas witb well. subshrubs, occasionaly provided with defined seasonality. · pseudobulbs: Leaves · conduplicate, subcoriaceous to fleshy. Inflorescence L Encyclia ~bbreviata (Scbltr.) Dressler, terminal, rarely lateral, a 1- to many-flowered Brittonia 13: 264: 1961. Epidendrum raceme, panicle or umbel. Flowers small to abbreviatum Schltr., Repert. Spec. Nov. Regni large and showy. Sepals and petals subsimilar, Veg: 3: 107. 1906. Epidendrum prorepens free, usually spreading; the lip simple or lobed, Ames, Sched. Orch. 2: 33. 1923. (Fig." 15). adnate to the column, usually with fleshy calli PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 985

smm

2.cm

1 cm 2 cm

Fig. 15. Encyclw abbreviata (Schltr.) Dressler. A- Habit. B - Flower. C- Column and lip, lateral view (sepals and petals removed). D- Dissected perianth. E- Colum, ventral view. F- Pollinaria and operculum. Illustration voucher:Luis Mena s.n. 986 REVISTA DE BIOLOGIA TROPICAL at the base. Column fused with the lip just to serruliferum Schltr., Repert. Spec. Nov. Regni the apex, often provided with a hooded Veg. Beíh. 19: 44. 1923. (Fíg. 17). clinandrium. Pollinia 4, waxy. Plant epiphytic, cespitose, without A genus of about 600 species, widespread pseudobulbs, with abbreviate, foliaceous in American tropics and subtropics from stems, up to 5 cm tall. Stems f!attened, to 5 cm southern USA to Argentina. long, 0.8 cm in diameter, completely concealed by closely appressed leaf sheaths, 5- l. Epidendrum amparoanum Schltr., to 11-leaved. Roots filiform, flexuous, Repert. Spec. Nov. Regni Veg. Beih. 19: 34. glabrous. Leaves distichous, elliptic-ovate, 1923. (Fig. 16). fleshy, retuse, dorsally apiculate, 1-2.3 cm Phmt epiphytic, cespitose, without long, 0.6-0.9 cm wide, not articulated with the pseudobulbs, with short rhizome and sheaths, the margíns somewhat revolute. foliaceous stems up to 35 cm tall. Stems Inflorescence terminal from the upper leaf, elongate, flattened, 35 cm long, to 1.4 cm in sessile, 1-2 flowered. Flowe:rs small, pale diameter, completely enfolded by the tubular, green wíth purplish suffusion, the lip emerald leaf-bearing sheaths, 5- to 9-leaved. Roots green. Sepals subsimilar, elliptic-lanceolate, fleshy, flexuous, glabrous. Leaves elliptic­ acute, the margins somewhat denticulate, oblong, coriaceous, retuse, 3.5-8 cm long, 1-3 dorsally carinate, the lateral sepals falcate, cm wide. Inflorescence terminal, a short concave, with a serrulate carine, 10-12 mm subumbellate, subsessile raceme bearing 3-5 long, 4 mm wide. Petals oblique, narrowly flowers, produced from the axil of the upper lanceolate, subacute, the apical margins leaf. Flowers medium sized, pale green. denticulate, 9 mm long, 2.8 mm wíde. Lip with Sepals subsimilar, lanceolate-elliptic, acute, a ligulate claw adnate to the column, the the lateral sepals somewhat falcate, 32 mm lamina ovate, somewhat cordate at the base, long, 10 mm wide. Petals linear elliptic, acute, acute, mucronate, 9 mm long, 5.5 mm wide. 28 mm long, 5 mm wide. Lip with a short claw Column short, stout, with a 3-toothed adnate to the column, the lamina 3-lobed, clinandrium, the margins erose, 5 mm long. widely reniform, deeply cordate at the base, Pollinia 4, laterally compressed, on a short somewhat 3-lobed at apex, apiculate, 28 mm caudicle. Anther cap hemiglobose, 4-celled. long, 35 mm wide; the disc with two Distribution: Endemic to Costa Rica. thickened, rounded, slightly diverging calli. Parque Nacional Manuel Antonio: Column clavate, with a terminal tooth, Widespread within the park, it is usually found margins of the clinandrium erose, 12 mm long. in lighted spots just to the seashore, closely Pollinia 4, laterally compressed, on a short appressed to the host tree but sometimes caudicle. Anther cap hemiglobose, 4-celled. loosely hanging from the branchlets. Plants Distribution: Costa Rica and Panama, exposed to full sunlight present a strong purple from sea leve} to about 1,500 meters. pygmentation on the leaves and the flowers. Parque Nacional Manuel Antonio: The The species is common at PNMA. species is widespread within the park, where it Phenology: flowering mostly occurs in is obiquitarious in all vegetation units just to April-May. the seashore. The flowers emit a plastic-like smell, that is stronger in the last hours of the 3. Epidendrum isomerum Schltr., Repert. day. Very common at PNMA. Spec. Nov. Regni Veg. 2: 132. 1906. Phenology: flowering mostly occurs in Epidendrum guayasense Schltr., Repert. Spec. July-August. Nov. Regni Veg. 8: 73. 1921. Plant epiphytic, pendent, without 2. Epidendrum congestum Rolfe, Kew pseudobulbs, up to 130 cm tal!. Stems Bull. 1913: 29. 1913. Epidendrum foliaceous, simple or commonly branched, PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 987

¡ ___ 2. cm 3 cm

Fig. 16. Epidendrum amparoanum Schltr. A - Habit. B - Flower. C - Column and lip, lateral view (sepals and petals removed). D - Dissected perianth. E- Column, ventral view. F- Pollínaria and operculum. Illustration voucher: F Pupulin 328. 988 REVISTA DE BIOLOGIA TROPICAL

1 cm

5 mm 2 mm

1 cm 1 cm

Fig. 17. Epidendrum congestum Rolfe. A- Habit. B- Flower. C- Column and lip, lateral view (sepals and petals removed). D - Dissected perianth. E - Colurim, ventral view. F - Pollinaria and operculum. Illustration voucher: F. Pupulin 296. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 989 flexuous, reaching 1 m or more in lenghth, 0.2 22. 1845. Epidendrum spruceanum Lindl., Fol. cm in diameter, the short branches about 10-12 Orch. Epidendrum 80. 1853. Epidendrum cm long, leafy toward the apex of new growths, nocturnum var. tridens Cogn. in Mart., Fl. Bras. covered with papyraceous sheaths at the base of 3 (5): 136. 1898. Epidendrum buenaventurae older stems. Roots fibrous, flexuous, glabrous, Lehm. & Krzl. in Engler, Bot. Jahrb. Syst. 26: produced at the base of the main stem. Leaves 470. 1899. Phaedrosanthus nocturnus Ktze. in linear, flat to subterete, acute, to about 1O cm Post & Ktze., Lex. Gen. Phanerog. 429. 1904. long, 0.2-0.4 cm wide, articulated to the Auliza nocturna Small, Fl. Miarni 56. 1913. sheaths, deciduous with age. Inflorescence Epidendrum leucarachne Schltr., Repert. Spec. terminal at the apex of lateral branches, 1- Nov. Regni Veg. Beih. 7: 136. 1920. flowered. Flowers small, yellowish green. Epidendrum tunguraguae Schltr., Repert. Spec. Sepals subsimilar, linear-lanceolate, acute, the Nov. Regni Veg. Beih. 8: 87. 1921. margins somewhat revolute, 11-15 mm long, Epidendrum oliganthum Schltr., Repert. Spec. 2.5-3 mm wide. Petals oblique, linear-elliptic, Nov. Regni Veg. Beih. 9: 90. 1921. a:cute, 11-13 mm long, 1.2-1.5 mm wide. Lip Epidendrum nocturnum var. panamense unguiculate, the claw adnate to the column, the Schltr., Repert. Spec. Nov. Regni Veg. Beih. 17: lamina narrowly lanceolate, canaliculate, acute, 36. 1922. Amphiglottis nocturna Britt., Sci. fleshy, 9-12 mm long, 4.5 mm wide when Surv. Porto Rico & Virgin Isl. 5 (2): 200. 1924. spread out; disc with a short, V-shaped callus (Fig. 18). and a median keel extending from the base to Plant epiphytic, erect to pendent, without the middle of the blade. Column short, stout, pseudobulbs, up to about 50 cm tall. Stems adnate to the lip just to the apex, with a foliaceous, to 45 cm long, 0.5 cm in diameter, cucullate clinandrium, the margins entire, 7 covered with tubular leaf sheaths. Roots mm long. Pollinia 4, laterally compressed, on a fleshy, flexuous, glabrous. Leaves elliptic­ short caudicle. Anther cap hemiglobose, 4- oblong, subcoriaceous, acute, 5-11 cm long, celled. 1.5-3 cm wide, articulated to the sheaths, Distribution: From Mexico to Panama, deciduous with age. Inflorestence terminal, a widespread in lowland wet forests. short, few- successively flowered raceme, Parque Nacional Manuel Antonio: enfolded by 2-3 papyraceous bracts at the base. Although 1 have not yet found plants of Flowers large, fragrant at night, sepals and Epidendrum isomerum within PNMA, petals green, the lip white, turning creamy Bolaños et al. (1983) recorded it in their list. yellow with age. Sepals subequal, narrowly The species is widespread and relatively linear, acute to acuminate, 8 cm long, 0.8 cm common outside the protected area and it may wide. Petals oblique, narrower than sepals, be found southward also in mangrove, where it acute, 7.5 cm long, 0.5 c,n wide. Lip has been collected from Rio Esquinas in Golfo unguiculate, the claw adnate to the column, the Dulce (R. Soto s.n., CR!). Rare at PNMA. lamina prominently 3-lobed, the midlobe Phenology: flowering probably in May­ acicular to subfiliform, the laterallobes broad, June. obliquely elliptic-ovate, obtuse to retuse, entire lip 6.5 cm long, 3 cm wide between the lateral 4. Epidendrum nocturnum Jacq., Enum. lobes; disc with a pair of short, slightly Syst. PL 29. 1760. Epidendrum cárolinianum diverging calli and a central keel running Lain., Encyél. ·1: 182. 1783 .. Nyctosma . toward the midlobe. Column clavate, stalkétl, noCturna Raf., Fl. Tellur. 2: 9. 1837. adnate to the lip just to the apex, with a Epidendrum tridens Poepp. & Endl., Nov. Gen. cucullate clinandrium, the margins erose, 22 et Sp. 2: 2. 1838. Epidendrum longicolle Lindl., mm long. Pollinia 4, laterally compressed, on Bot. reg. 24: mise. 34. 1838. Epidendrum a short caudicle.,,. Anther cap subrhombic, discolor Rich. & Gal., Ann. Sci. Nat. ser. 3, 3: cucullate, 4-celled. 990 REVISTA DE BIOLOGIA TROPICAL

Fig. 18. Epidendrum nocturnum Jacq. A- Habit. B- Column and lip, lateral view (sepals and petals removed). C- Dissected perianth. D - Column, ventral view. E - Pollinaria and operculum. Illustration voucher: F Pupulin 335. PUPULIN: Orchid Aorula of Parque Nacional Manuel Antonio 991

Distribution: From Florida to tropical spread out. Column short, stout, adnate to the South America and the West Indies, lip near to the apex, with a cucullate, 3-toothed widespread at low elevations. clinandrium, 4 mm long. Pollinia 4, on a short Parque Nacional Manuel Antonio: The caudicle. Anther cap cucullate, 4-celled. species is widespread within the protected Distribution: From Honduras to Ecuador area, where it may be found in lighted and the Guyanas, local in lowland forests. situations in secondary forest and on short Parque Nacional Manuel Antonio: The trees along the sea. It was once much more record is based on a single collection from a frequent on Terminalia catappa and Tabebuia high, main branch of Pithecellobium saman rosea trees on the beach at the entran ce of the along a grassland and no other observations park, but in recent years these populations were made in the neighbouring areas. The have been severely reduced for unknown species is rare at PNMA. reasons. The species should now be regarded Phenology: flowering occurs from as occasional at PNMA. December to February. Phenology: flowering occurs sporadically throughout the year. 6. Epidendrum stamfordianum Batem., Orch. Mex. & Guat. t. 11. 1838. Epidendrum 5. Epidendrum sculptum Rchb. f., basilare Kl., Allg. Gartenz. 11: 193. 1843. Bonplandia 2: 89. 1854. Epidendrum Epidendrum cycnostalix Rchb. f., Bot. Zeit. colonense Ames, Sched. Orch. 1: 14. 1922. 10: 731. 1852. Epidendrum stamfordianum (Fig. 19). var. pictum Lemaire, Jard. Fl. 3: t. 251. 1852. Plant epiphytic, pendent, without Epidendrum staínfordianum var. parviflorum pseudobulbs, up to 80 cm talL Stems Regel, Ind. Sem. Hort. Petro. 19. 1856. foliaceous, simple or 'commonly' many­ Epidendrum stamfordianum var. wallacei branched, sómewhat flexuous, about 0.5-0.8 Rchb.f., Gard. Chron. ser. 3, 1: 543. 1887. cm in diameter, densely leaved, covered with Epidendrum stamfqfdianum var. leeanum papyraceous sheaths at the base of older stems. Rchb.f., Gard. Chron. ser. 3, 3: 521. 1888. Roots fibrous, flexuous, glabrous, produced at Epidendrum stamfordianum var. the base of the main stem, about 1 mm in lawrenceanum hort. ex Stein, Orchideenb. 240. diameter. Leaves elliptic-oblong, fleshy, 1892. Psilantheum basilare Kl. ex Stein., obtuse .to retuse, 2-4.9 cm long, 0.5-1.2 cm Orchideenb. 238. 1892. Auliza stamfordianum wide, articulated to the sheaths, deciduous .(Batem.) Brieg., Die Orchideen 3. Aufl. 1 (9): with age. Intlorescence terminal at tl).e apex of 548. 1977. (Fig. 20). lateral branches, 1-flowered or commonly 2- Plant epiphytic, cespitose, erect, to about flowered. Flowers small, yellowish green, 50 cm tall. Roots fleshy, flexuous, glabrous. fleshy, scented. Sepals subsimilar, lanceolate Pseudbulbs stalked, fusiform, enfolded by to linear-oblong, obtuse, the margins many large bracts becoming papyraceous with somewhat revolute; 9-10.5 mm long, 2.5-3 mm age, 8-28 cm long, 1.5-2.2 cm in diameter, 2- wide. Petals oblique, linear-oblong, obtuse, 10 to 4-leaved near the apex. Leaves rather mm long, 2 mm wide. Lip unguiculate, the variable, oblong-elliptic to ligulate or claw adnate to the column, the lamina 3-lobed, oblanceolate, obtuse, coriaceous, 10-22 cm ovate-lanceolate, cordate at the base; the long, 2.5-6.5 cm wide. Intlorescence lateral lateral lobes elliptic, erect, reaching about the from a short shoot along the rhizome, an erect middle of the lip; midlobe lanceolate, to slightly arcuate, simple or rarely paniculate apiculate; disc with a pair of short, mamillate raceme, usually many-flowered raceme to 50 calli at the base of the lateral lo bes and two low cm long, enfolded by 2-3 triangular bracts. ridges extertding to the middle of the midlobe; Flowers medium sized, showy, ' sepals and entire lip 11 mm long, 5.5 mm wide when petals greenish yellow marked with dark 992 REVISTA DE BIOLOGIA TROPICAL

20 cm

1 cm Z.cm

Fig. 19. Epidendrum sculptum Rchb.f. A- Habit. B - Flower. C- Coluum and lip, lateral view (sepals and petals removed). D- Dissected perianth. E- Coluum, ventral view. F- Pollinaria and operculum. Illustration voucher: F. Pupulin 373. PUPULIN: Orchíd Florula of Parque Nacional Manuel Antonio 993

3 cm 5mm

F

.s mm

3 cm

Fig. 20. Epidendrum stamfordianum Batem. A - Habit. B - Flower. C - Column and lip, lateral view (sepals and petals removed). D - Dissected perianth .. E - Column, ventral view. F - Operculum and pollinaria. Illustration voucher: F. Pupulin 370. 994 REVISTA DE BIOLOGIA TROPICAL

purple, the lip white variously spoted or suffuse base of the column, the lamina exceeding the with lilac to purple. Sepals subequal, elliptic­ sepals. Column erect, nearly terete, footless, oblanceolate, acute, the lateral sepals falcate, sometimes provided with rudimentary wings. concave near the apex, 1.8-2 cm long, 0.5 cm Pollinia 2, waxy. wide. Petals oblique, linear-oblong, acute, 1.8 A neotropical genus of 3 to 5 species, cm long, 0.3 cm wide. Lip unguiculate, the often restricted to the twigs of the host trees. claw adnate to the column, the lamina prominently 3-lobed, the midlobe transversely l. lonopsis satyrioides (Sw.) Rchb.f., oblong, with a short claw at the base, crenulate Walp. Ann. Bot. Syst. 6: 683. 1861. to serrulate; the laterallobes broad, subqadrate­ Epidendrum satyrioides Sw., Prod. 123. 1788. elliptic, rounded, crenulate; disc provided with Dendrobium testiculatum Sw., Nov. Act. a short bilobed callus at the base and a median, Upsal. 6: 83, 1799. lonops'is testiculata (Sw.) low keel running to Ihe middle of the midlobe; Lindl., Gen. & Sp. Orch. Pl. 193. 1833. the entire lip 1.7 cm long, 2 cm wide between lonopsis teres Líndl., Bot. Reg. 24, mise. 95. the lateral lobes. Coiumn clavate, stalked, 1838. (Fig. 21 ). adnate to the lip just to the apex, with a Plant epiphytic, mainly on twigs and cucullate, 3-toothed clinandrium, 9 mm long. small branches, to about 15 cm tall, with PoUinia 4, lateral! y compressed, in two pairs of abbreviate rhizome. Roots fibrous, flexuous. different size, on a short caudicle. Anther cap Pseudobulbs inconspicuous, elliptic to cucullate, lanceolate, 4-celled. subcylindric, laterally flattened, to 4 mm long, Distribution: Widespread at low about 2 cm wide, nearly completely hidden by elevations in Central and northern South 2-5 leafy sheaths, monophyllous. Leaves America, from Mexico to Panama, Colombia terete, linear to linear-attenuate, acute, to 10- and Venezuela. 12 cm long, about 0.3 cm in diameter. Parque Nacional Manuel Antonio: Within Inflorescence a lateral, successively flowered the park the species is mainly restricted to the raceme, to 16 cm long (but usually shorter). main branches and trunks of re!ictual, large Flowers small, white with obscure rose Pithecellobium spp. trees in pastures. Here it is stripes. Sepals campanulate, not completely not uncommon to find hugues specimens with spreading. Dorsal sepal lanceolate-elliptic, more than 20 large, leafy pseudobulbs, each acute, to 9 mm long, 3-4 mm wide. l .. ateral one exceeding 50 cm in length. Frequent at sepals elliptic-lanceolate, acute, connate at the PNMA. base for about one third of their lenghth, to lO Phenology: flowering occurs in March­ mm long, 3.5 mm wide. Petals ovate-elliptic, April. obtuse, minute! y mucronate, lO mm long, 3.5 mm wide. Lip clawed, cuneate, obtuse to retuse, slightly concave toward the apex, with 11. IONOPSIS Kunth two rudimentary auricles near the base, the disc provided with two diverging, thick keels; Small, cespitose, epiphytic herbs with entire lip 11 mm long, 5 mm wide. Column inconspicuous stems, 1- 2-foliate at apex, short, with two obscure wings, to 3 mm long. subtended by foliaceous sheaths. Leaves Pollinia 2, subspherical, on a ligulate stipe; conduplicate, coriaceous to subterete. viscidium elliptic. Anther cap hemiglobose, Inflorescence lateral, a few- to many-flowered, 2-celled. erect to archíng raceme or panicle. Flowers Distribution: From Costa Rica to northern small, whíte to pale lilac somewhat striped South America, widespread in wet forests and with rose-purple. Sepals and petals spreading; orchards. the lateral sepals connate producing at the base Parque Nacional Manuel Antonio: Plants a short sac; lip simple, clawed, adnate to the of /onopsis satyrioides have so far escaped - PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 995

3mm .1 cm F

1 cm

3 cm

Fig. 21./onopsis satyrioides (Sw.) Rchb.f. A- Habit. 8 - Flower. C - Colurnn and lip, three quarter view (sepals and petals removed). D - Dissected perianth. E - Colurnn, ventral view. F - Pollinarium and operculum. Illustration voucher: F. Pupulin 309. 996 REVISTA DE BIOLOGIA TROPICAL attention at PNMA and my collection is from green vegetative apex. Pseudobulbs ovate to the low hills northern to the park. However, the elliptic, laterally flattened, to 3.2 cm long, species is a common inhabitant of disturbed about 1.5 cm wide, basally covered by 2-3 habitats and orchards (mainly on Citrus and Jeafy sheaths, monophyllous. Leaf Psidium spp.) and its presence within the conduplicate, linear to linear-elliptic, obtuse, protected area is likely. It should be considered slightly emarginate, to 10-12 cm long, 1.2-2 rare at PNMA. cm wide. Inflorescence basal, to 16 cm long, Phenology: flowering occurs from 1- to many-flowered (mainly 3-flowered); September to December. peduncle terete; bracts triangular-lanceolate, acute, about 0.7 cm long. Flowers small, greenish yellow spotted with purple red. 12. LEOCHILUS Knowles and Westcott Dorsal sepal elliptic-obovate, acute, to 5.5 mm long, 0.3-0.4 mm wide. Lateral sepals Small, cespitose, epiphytic herbs with obliquely elliptic-obovate, acute to slightly ovoíd, laterally compressed pseudobulbs, 1- 2- apiculate, to 7 mm long, 0.3 mm wide. Petals foliate at apex, subtended by 1-2 foliaceous linear, apiculate, to 6-7 mm long, about 3 mm sheaths. Leaves conduplicate, elliptic to wide. Lip obovate, obtuse to retuse, flat, with oblong, subcoriaceous. Inflorescence lateral, a a small, concave nectary at the base provided few- to many- successively flowered raceme with pilose margins; callus trapezoidal, with a or panicle. Flowers small, greenish-yellow central groove, papillose; entire lip 9 mm long, usually stripped with brown. Sepals and petals 5 mm wide. Column short, with a pair of spreading; the lateral sepals variously connate; triangular, acute stigmatic arms, to 2.5 mm lip simple or 3-lobed, provided near the base long. Pollinia 2, subspherical, on a short, with a fleshy callus serving as an open nectary ligulate stípe; viscidium peltate, brown. filled with oil. Column erect, nearly terete, Anther cap hemiglobose, 2-celled. footless, often provided with stigmatic arms. Distribution: From Mexico to Panama and Pollinia 2, waxy. northern South America. Common at lower A neotropical genus of about 9 species, elevations on Pacific coasts in Central sometimes restricted to the twigs of the host America. trees. Parque Nacional Manuel Antonio: Epiphytic on Croton sp. at the edge of l. Leochilus scriptus (Scheidw.) Rchb.f., secondary forest. Plants of L. scriptus were Xenia Orchid. 1(1 ): 15, 1854. Clyptosanus found growing in medium-shaded conditions, scriptus Scheidw., Allg. Gartenzeit. 11: 101, on little branches and twigs without any 1843. Leochilus herbaceus Lindl., Bot. Reg. mosses. The species is rare at PNMA. 30: Mise. 90, 1844. Cryptosaccus scriptus Phenology: flowering mainly occurs from Scheidw. ex Rchb.f., Xenia Orchid. 1(1): 15, December through February and March, but 1854. Oncidium herbaceum (Lindl.) Rchb.f., plants may flower also in other months. Ann. Bot. Syst. 6: 772, 1863. Oncidium scriptum (Scheidw.) Rchb.f., Ann. Bot. Syst. 6: 772, 1863. Leochilus major Schltr., Repert. 13. LOCKHARTIA Hook. Spec. Nov. Regni Veg. Beih. 15: 209, 1919. Leochilus powellii Schltr., Repert. Spec. Nov. Cespitose, epiphytic herbs without Regni Veg. Beih. 17: 81, 1922. Leochilus pseudobulbs, the foliaceous stems erect or retusus Schltr., Repert. Spec. Nov. Regni Veg. pendent, completely envolved by the Beih. 19: 256, 1923. (Fig. 22). distichously imbricating leaves. Leaves short, Plant epiphytic, to about 15 cm tall, with laterally flattened, triangular, coriaceous. abbreviate rhizome. Roots fleshy, white with Inflorescence lateral, few- to many- PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 997

3 mm

1 cm __J '------

1 cm <'\cm

Fíg. 22. Leochilus scriptus (Scheidw) Rchb. f. A- Habit. B- Juvenile plant. C- Flower. D- Dissected perianth. E- Column and lip, lateral view (sepals and petals removed). F- Pollinarium and operculum. Illustration voucher: F. Pupulin 374. 998 REVISTA DE BIOLOGIA TROPICAL successively flowered raceme or panicle, lobes subtriangular, rounded, 0.5 mm long, 1 emerging from the axils of the leaves, the mm wide; the midlobe widely obovate to pedicel often covered with conspicuous subrhombic, emarginate, with a short, rounded membranaceous bracts. Flowers small, yellow apicule; the disk with a trapeziform callus, or white, usually blotched with reddish brown distally bilobed, provided at the base with a on the lip. Sepals subequal, free, spreading or tuft of distinct papillae, and three irregular sometimes reflexed; petals usually broader; lip rows of rounded, flat tubercles in front of the simple or 3-lobed, the laterallobes rounded to callus running just below the half of the lip. lnear, the midlobe variously 2- to 4-lobed, Column short, with two triangular­ provided near the base with a complex callus. dolabriform wings, 1.3 mm long, 1.9 mm Column short, footless, provided with large wide. Anther cap cucullate, papillose, 2- apical wings. Pollinia 2, waxy. celled. Pollinia 2, pyriform-obovate, on two A neotropical genus of about 30 species, linear stipes; viscidium subguadrate, white. usually found on trunks and large branches of Paratypes: Costa Rica. Prov. Puntarenas: trees. Aguirre, Parque Nacional Manuel Antonio, F. Pupulin 324 (SEL, Herb. Pupulin). l. Lockhartia pandurata Pupulin sp. nov. Etymology: Narned in reference to the Species Lockhartiae micranthae Rchb.f. pandurate lamina of the lip. similis, sed lobis lateralibus labelli Distribution: Known only from Costa subtriangularibus rotundatis multo brevíoribus Rican central Pacific lowlands. et caespite tomentoso in disco labelli differt. Parque Nacional Manuel Antonio: Type: Costa Rica. Prov. Puntarenas: Aguirre, Epiphytic on lower canopy and trunks; just to Parque Nacional Manuel Antonio, in disturbed the seashore on the high branches of tall trees. primary forest, 80 ni, 11 July 1995, F. Pupulin Common at PNMA. 313 (holotype,USJ; isotype, CR). (Fig. 23). Phenology: flowering occurs m Herb epiphytic, cespitose, without December-January. pseudobulbs, to about 25 cm tall. Roots Lockhartia pandurata is closely allied to filiform, 0.8 mm in diameter. Stems L. micrantha, with which it has been confused foliaceous, erect, becoming pendent with age, in herbarium specimens. In his key to Costa 10 to 25 cm long, 1-1.5 cm wide. Leaves Rican and Panamian Lockhartia species, distichous, imbricate, triangular in lateral view, Dressler (1993) divíded the genus in two apically obtuse to rounded, 1.4-1.5 cm long, groups primarily based on the presence or lateral width 0.6-0.7 cm. Inflorescence a absence of definite lateral lobes of the lip, but terminal or subterminal raceme of few flowers, L. pandurata fills the gap between the two 1.3-1.5 cm long, provided with 1-2 triangular­ groups. In L. pandurata the lateral lobes are ovale, acute bracts, 2 mm long, 1.6 mm wide; reduced to very short, subtriangular, rounded the ovary with pedicel about 8 mm long; floral ear-like projections. bracts suborbicular, acute, 5.3 mm long, 3.8 mm wide. Flowers small for the genus, 6 mm in width, pale yellow with pale red-brown 14. MAXILLARIA Ruiz & Pav. spots on Iip and column. Dorsal sepal suborbicular, apiculate, concave, 3 mm long, Epiphytic or lithophytic herbs with short 2.5 mm wide. Lateral sepals ovate-elliptic, to elongate, erect to pendent stems, with or apiculate, concave, 3.2 mm long, 2 mm wide. without clustered to distant pseudobulbs, 1- 3- Petals elliptic, slightly concave near the apex, foliate at apex. Leaves conduplicate, thin to 3.4 mm long, 2.2 mm wide. Lip 3-lobate, coriaceous, persistent. Inflorescence a scape pandurate, 3.5 mm long, 3.5 mm wide (2.8 mm from the base of pseudobulbs or lateral from wide between the lateral lobes); the lateral leaf axils or from the developing new growth, PUPULIN: Orchid Aorula of Parque Nacional Manuel Antonio 999

5 mm

E

2 mm

5 tnm 5cm

Fig. 23. Lockhartia pandurata Pupulin. A - Habit. B - Aower. C - Dissected perianth. D - Column, ventral view. E - Pollínarium and operculum. Illustration voucher: F. Pupulin 313. 1000 REVISTA DE BIOLOGIA TROPICAL

1 to many. Flowers small to large. Sepals and linear-oblanceolate, acute, curved, 13 mm petals spreading; the lateral sepals sometimes long, 4 mm wide. Lip obscurely 3-lobed, connate, often forming short mentum with the elliptic-lanceolate, contracted at the base and column base; lip simple or 3-lobed, articulated articulate with the column foot, to 16 mm long, with or adnate to the column, the laterallobes 9 mm wide when spread out; lateral lobes erect, the midlobe spreading or reflexed, rounded, erect; midlobe subobtuse, thickened usually provided at the base with a fleshy and farinose to minutely papillose; disk with a callus. Column erect, nearly terete, with or ligulate, fleshy, thickened, farinose callus. without a conspicuous foot, rarely winged. Column elongate, arcuate, semiterete, to 12 Pollinia 4, waxy. mm long, produced at the base into a short A neotropical genus of sorne 600 mainly foot. Pollinia 4, in two different sized pairs, on poorly known species, widespread from a short, rounded, basally dilated stipe. Anther Mexico to Bolivia and Brazil, and the West cap cucullate, glabrous, 1-celled. Indies. Distribution: A widespread and rather common species ranging from Florida, l. Maxillaria crassifolia (Limll.) Rchb.f., Mexico, through Central America, to Panama, Bonpl. 2: 16, 1854. Heterotaxis crassifolia the Caribbeam Islands, Venezuela, Brazil and Lindl., Bot. Reg. 12: t. 1028, 1826. Dicrypta Colombia. baueri Lindl., Gen. et Spec. Orch. PI, 44, 1830. Parque Nacional Manuel Antonio: Dicrypta crassifolia Lindl. ex Loud. Hort. Brit. Epiphytic on Hymenaea courbaril (guapinol) Suppl. 3: 536, 1839. Epidendrum sessile and Cynometra hemitomophylla (guapinol Swartz, Prodr. Regn. Veg. Ind. Occ. 122: 1878, negro) trees mainly in primary or mature non Maxillaria sessilis Lindl. Maxillaria secondary forest. Plants are generally found on sessilis Fawc. & Rendle, Fl. Jam. 1: 120, 1910, large trunks in medium shaded conditions. non Lindl. Maxillaria gatunensis Schltr., Fedde Occasional at PNMA. Rep. Sp. Nov. Beih. 17: 68, 1922. (Fig. 24). Phenology: flowering occurs throughout Plant epiphytic, cespitose, pendent, with a the year. short rhizome, to about 35 cm tal!. Roots filiform, glabrous. Pseudobulbs oblong, 2. Maxillaria neglecta (Schltr.) L.O. inconspicuous, monophyllous, subtended by 2- Williams, Ann. Missouri Bot. Gard. 29: 348. 3 distichously imbricating, fleshy, foliaceous 1942. Omithidium anceps Rchb.f., Beitr. Orch. sheaths, to 3 cm long, about 1.5 cm wide. Centr.-Amer. 75. 1866, non Maxillaria anceps Leaves coriaceous, linear to linear-oblong, Ames & C. Schweinf. Ornithidium neglectum obtuse to subacute, forming at the base a more Schltr., Fedde Rep. Sp. Nov. Beih. 19: 242, or less complanate petiole, 7-35 cm long, 1.4- 1923. (Fig. 25). 2.8 cm wide. Inflorescence produced from the Plant epiphytic, pendent, with elongate, axils of the upper lea ves, generally 1-flowered, sometimes branching rhizome, to 50 cm tall. with a short peduncle 1 to 2.2 cm long, Roots filiform, glabrous. Pseudobulhs ligular­ provided by a scarious bract. Ovaries elliptic to widely elliptic, ancipitous, usually pedicellate, subclavate, about 1 cm long. very distant, monophyllous, 1.5-3 cm long, Flowers of moderate size, pale yellow, the lip about 0.7-1.5 cm wide, the long internodes marked with purple. Dorsal sepal oblong­ enveloped in closely imbricating, papery elliptic to lanceolate, subacute, conspicuously bracts. Leaves subcoriaceous, ligular, acute, concave, fleshy, 17 mm long, 6.5-7 mm wide. 5.5-12 cm long, 1.3-1.9 cm wide, forming at Lateral sepals obliquely ovate-oblong, the base a short petiole. Inflorescences subacute, dorsally carinate, 16 mm long, 6 mm produced from the base of the pseudobulb, 1- wide, adnate to the column, forming an flowered, short scapes produced in dense inconspicuous, rounded mentum. Petals fascicles, about 1 cm long, partially hidden by PUPULIN: Orchid Florula ofParque Nacional Manuel Antonio 1001

1 cm

5 rnm

Jmm

2cm

Fig. 24. Maxillaria crassifolia (Lindl.) Rchb.f. A - Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E- Column, ventral view. F- Column, ventral view, anther removed. G- Pollinarium and operculum .. Illustration voucher: F. Pupulin 312. 1002 REVISTA DE BIOLOGIA TROPICAL

F

5 mm 3 mm 1 cm

Fig. 25. Maxillaria neglecta (Schltr.) L. O. Williams. A- Habit. B - Flower. C- Dissected peranth. D- Flower, lateral view. E- Column and lip, lateral view. F- Column, ventral view. G - Pollínarium and operculum. lllustration voucher: F. Pupulin 363. PUPULIN: Orchid Flonlla of Parque Nacional Manuel Antonio 1003

short glumaceous bracts. Flowers small, red lateral lobes. Sepals subsimilar, white, the lip with yellow mídlobe. Dorsal lanceolate to ovate-lanceolate, acute, sepa! elliptic-ovate, subacute, conspicuously concave toward the base, the lateral sepals concave, fleshy, 6 mm long, 3 mm wide. adnate to the column foot to form a short Lateral sepals ovate, somewhat falcate mentum, 13-15 mm long, 5 mm wide. Petals subacute, dorsally carinate, connate at the base lanceolate, acute, erect, 12 mm long, 3 mm to produce a rounded mentum, 6-7 mm long, 4 wide. Lip 3-lobed, articulated with the mm wide. Petals oblong-ligular, subacute, 5-6 column foot, oblong-ligular, the lateral lobes mm long, about 2 mm wide. Lip unguiculate, erect, rounded, the midlobe ovate, subacute, 3-lobed, the claw continuous with the column thickened, minutely puberulent 12 mm long, foot, then geniculate, to 6 mm long, 4 mm 5 mm wide across the laterallobes; disc with wide when spread out; lateral lobes obliquely a linear, fleshy callus. Column semiterete, triangular, erect; midlobe fleshy, widely ovate­ curved, dilated at apex, the base produced subcordate, obtuse; disk with a ligulate, low into a foot, about 5 mm long. Anther cap callus. Colmnn short, semiterete, to 2 mm cucullate, pubescent. Poninia 4 on a saddle­ long, produced at the base into a conspicuous shaped stipe. foot. Pollinia 4, in two different sized pairs, on Dístribution: A widespread but not a short, dilated stipe. Anther cap cucullate, common species in lowlands and middle glabrous, 2-celled. elevation forests from Nicaragua to Panama. Distribution: A widespread and locally Parque Nacional Manuel Antonio: The common species rang1ng from Guatemala to single collection from the protected area is Panama in lowland wet forests. from the low branches of a tree at the edge of Parque Nacional Manuel Antonio: primary forest, where the plants were growing Epiphytic on Cynometra hemitomophylla in shaded conditions. Rare at PNMA. (guapinol negro) and other tall trees, mainly in Phenology: Flowering recorded in primary and mature secondary forest. Plants February to April. are generally found on lower branches in medium shaded conditions. Occasional at 4. Maxillaria ponerantha Rchb.f., PNMA. Bonplandia 2: 17. 1854. (Fig. 26). Phenology: flowering mainly occurs from Plant epiphytic, small, with ascending, July to September. freely branching stems, to 15 cm tall. Roots filiform, glabrous, produced only at the base of 3. Maxillaria oreocharis Schltr., Fedde Rep. the main stem. Pseudobulbs elliptic to ovate­ Spec. Nov. Beih.: 17: 69. 1922. elliptic, anctpltous, l-2 cm apart, Plant epiphytic, with scandent rhizome, monophyllous, 1-1.3 cm long, about 0.8 cm erect or pendent, about 15 cm tall. Roots wide, subtended by 2-4 cataphylls, the leaves slender, flexuous, mainly produced from the articulated with the sheaths and ultimately rhizome under the basal pseudobulbs. deciduous, the intemodes enveloped in loosely Psendobulbs elliptic to ovate-elliptic, imbricating, papery bracts. Leaves linear­ complanate, green, to 3.2 cm long, 1.5-2 cm oblong, obtuse, unequally bilobed, 1.3-2.8 cm wide, covered with papyraceous sheaths, long, 0.4-0.8 cm wide. Inflorescences monophyllous. Le~wes linear, acute to produced from the axil of cataphylls, 1- acuminate, contracted at the base into a very flowered, single scapes about 5 mm long. short petiole, to about 20 cm long, 0.8-1 cm Flowers small, very dark purple. Dorsal sepai wide. Inflorescence basal, from the axils of ovate-lanceolate, subacute, concave, fleshy, 6- the upper sheaths, a single-flowered raceme 7 mm long, 2.8 mm wide. Lateral sepals to 3.5 cm long. Flowers spreading, white elliptic-lanceolate, somewhat falcate, acute, marked with red and the lip yellow with dark dorsally carinate, free, 7 mm long, about 3 mm 1004 REVISTA DE BIOLOGIA TROPICAL

E F ~ 3 cm 2 mm

5 mm

5 mm 1 cm

Fig. 26. Maxillaria ponerantha Rchb.f. A- Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral view. F- Pollinarium and operculum. lllustration voucher: F Pupulin 345. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1005 wide. Petals elliptic-oblong, acute, 5 mm long, Plant epiphytic, erect, with abbreviate 1.3 mm wide. Lip narrowly ovate, obscurely rhizome, to 15 cm tall. Roots filiform, 3-lobed, 6 mm long, 3 mm wide; lateral lobes glabrous, produced at the base of pseudobulbs. elliptic, rounded, erect; midlobe fleshy, Pseudobulbs ovate-pyriform, sometimes subquadrate, retuse; disk with a ligulate, fleshy stalked, ancipitous, sulcate, monophyllous, callus. Column elongate, semiterete, 5 mm 1.8-2.3 cm long, about 1 cm wide, subtended long, produced at the base into a short foot. by 2-5 papyraceous sheaths. Leaves linear to Pollinia 4, in two different sized pairs, on a oblong-lanceolate, unequally bilobed at apex, short, saddle-shaped stipe. Anther cap subcoriaceous, 7-12 cm long, 0.6-1 cm wide. cucullate, verruculose, 1-celled. Inflorescence terminal, often produced before Distribution: Widespread but not common the pseudobulb reachs maturity, a few­ from Costa Rica to northern South America. flowered raceme shorter than leaves, to 8-10 Parque Nacional Manuel Antonio: cm long including the peduncle, bearing 1-2 Epiphytic on trunks and large branches of trees acute, spreading bracts. Flowers small, in primary and disturbed primary forest. Plants fragrant, creamy white, marked with yellow at are only found on old trees with living the base of the lip. Sepals lanceolate, acute, bryophyte, in shaded conditions. Frequent at somewhat concave towards the apex, 10 mm PNMA. The taxon is accepted here as M. long, 2-2.5 mm wide; lateral sepals slightly ponerantha, but the consistent difference in falcate. Petals oblong-lanceolate, acute to color and minor morphological details in Costa acuminate, 8 mm long, 1.8 mm wide. Lip Rican populations may perhaps deserve narrowly ovate-ellíptic to linear, recurved near taxonomic recognition. the base , 6.5 mm long, 1.8 mm wide, smooth. Phenology: flowering mainly occurs in Column elongate, subclavate, arcuate, 4 mm July-August. long, contracted at the base. Pollinia 4, in two different sized pairs, on two short caudicles. Anther cap cucullate, carinate, 4-celled. 15. NIDEMA Britt. & Millsp. Distribution: A widespread species from southern Central America to Venezuela, Cespitose, epiphytic, herbs with erect Colombia, Peru, and the West Indies. stems, wíth clustered pseudobulbs, 1-foliate at Parque Nacional Manuel Antonio: Plants apex. Leaves conduplicate, linear, of Nidema ottonis have been found in primary subcoriaceous, persistent. Inflorescence a few­ and secondary forest, epiphyte on branches flowered raceme from the apex of and trunks in sunlighted spots, where they pseudobulbs. Flowers small. Sepals and petals form large populations. The species is frequent free, not completely spreading, lanceolate atPNMA. elliptic; lip entire, articulated with the column Phenology: Flowering mostly occurs from base, smooth. Column arcuate, semiterete, March to June. with obscure wings. Pollinia 4, in two pairs of different size. A neotropical genus of sorne 2 species, 16. NOTYLIA Lindl. widespread from Mexico to northem South America, and the West Indies. Cespitose, epiphytic herbs with clustered pseudobulbs, 1-foliate at apex, with or without l. Nidema ottonis (Rchb.f.) Britt. & conspicuous cataphylls. Leaves conduplicate, Millsp., Bahama Fl. 94. 1920. Epidendrum coriaceous, persistent. Inflorescence apendent ottonis Rchb.f., Hamb. Gartenzeit. 14: 213. 1858. raceme from the axil of upper cataphylls, Nidema boothii var. triandrum Schltr; Fedde Rep. many-flowered. Flowers small. Sepals and Sp. Nov. Beih. 17: 43. 1922. (Fig. 27). petals . spreadíng, the lateral sepals free to 1006 REVISTA DE BIOLOGIA TROPICAL

1cm

E F

2 mm

3mm

1cm

Fig. 27. Nidema ottonis (Rchb.f.) Britton & Millsp. A- Habit. B - Flower. C- Dissected perianth. D- Column and lip, lateral view. E- Column, ventral view. F- Pollinaria and operculum. Illustration voucher: F. Pupulin 318. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1007

variously connate; lip clawed, entire or obscurely shaded conditions. Notylia pittieri is probably lobed, articulated with the column base, smooth pollinated by bees of the genus Euglossa. The or with a keeled callus. Column erect, semiterete, species is occasional at PNMA. with dorsal anther. Pollinia 2, waxy. Phenology: flowering recorded in July­ A neotropical genus of sorne 45 species, August. widespread from Mexico to South America, and the West Indies. 17. ONCIDIUM Sw. l. Notylia pittieri Schltr., Beih. Bot. Centralbl. 36, Abt. 2: 502. 1918. (Fig. 28). Cespitose to scandent, epiphytic, rarely Plant epiphytic, pendent, to 20 cm tall, terrestrial herbs with foliaceous stem with very short rhizome. Roots fleshy, frequently thickened into definite · glabrous. Pseudobulbs clustered, ellipsoid, pseudobulbs, 1- to 2-foliate (rarely more) at _compress, 2.5 cm long, 0.7 cm wide, apex, with or without conspicuous cataphylls. monophyllous, subtended by leafy and non­ Leaves conduplicate or terete, subcoriaceous leafy scarious sheaths. Leaf coriaceous, to fleshy, persistent. lnflorescence a often abaxially finely rugulose, oblong-ligulate to flexuous, erect to pendent raceme, frequently elliptic, inequally bilobed at apex, obscurely paniculate, from the lateral bases of mucronate, to 10.2 cm long, 3.9 cm wide. pseudobulbs, usually many-flowered. Flowers Intlorescence basal, emerging from the axil of usually large and conspicuous, often yellow imbricating leaf base, pendent, racemose or and brown. Sepals and petals usually secundarily paniculate, to 19 cm long, many­ spreading, the lateral sepals free to variously flowered (to 80 and more); bracts small, connate; lip clawed, usually 3-lobed, often triangular-lanceolate, acuminate, 2 mm long. pandurate, the disc usually cristate or Flowers small, with greenish white sepals, tuberculate. Column erect, semiterete, often white petals with 3 yellow blotches, and white winged, provided at the base with a tabula lip. Dorsal sepal oblong-ligulate, obtuse, infrastigmatica, with terminal anther. Pollinia slightly acuminate, deeply revolute, concave, 2, usually deeply sulcate, waxy. 4.8 mm long, 1.5 mm wide. Lateral sepals A polimorphic genus of sorne 400 species, connate just to 1 mm from apex, broadly widespread in the Neotropics from Florida to oblong to obovate, the apex obtuse, reflexed, the West Indies and to Bolivia, Brazil and 4.1 mm long, 2 mm wide. Petals obliquely Argentina in South America. ligulate-subfalcate, acute, 4.2 mm long, 1.2 mm wide. Lip shortly clawed; lamina l. Oncidium ascendens Lindl., Bot.· Reg. triangular-subhastate, subobtuse, dorsally 28: sub. t. 4. 1842. Oncidium helicanthum carinate at apex, with a basal thickened, linear­ Krzl. in Engler, Pflanzenr. 4. 50, Heft. 80: 95. oblong callus produced by the extension of the 1922. Oncidium subulifolium Schltr., Fedde thick claw, 3.4 mm long, 1.9 mm wide. Rep. Sp. Nov. 10: 79. 1922. (Fig. 29). Column terete, slender, with dorsal anther, 2.5 Plant epiphytic, pendent, to 70 cm tall, mm long. PoUinia 2, obtriangular, laterally with very short rhizome. Roots fleshy, flattened, on a long, obtriangular attenuate glabrous, branched. Pseudobulbs clustered, stipe; viscidium elliptic, brown. Anther cap inconspicuous, rounded, to 1 cm long, 0.8 cm oblong, cucullate, 1.4 mm lol)g. wide, monophyllous, subtended by Distribution: Endemic to Costa Rica. papyraceous sheaths. Leaf fleshy, terete, Parque Nacional Manuel Antonio: Plants attenuate tow&rd the apex, adaxially were found as epiphyte on Citrus sp. (limón) canaliculate, to 70 cm long, 1.6 cm wide. and Cynometra hemitomophylla (guapinol Inflorescence lateral, emerging from the axil negro) in young secondary forest and in of the imbricating sheaths, paniculate, pendent, 1008 REVISTA DE BIOLOGIA TROPICAL

E

1mm

B

5 mm

5mm

Fig. 28. Notylia pittieri Schltr. A - Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Pollinarium and operculum. Illustration voucher: F. Pupulin 292. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1009

E

1cm 5mm A

F

a cm 3mm

Fig. 29. Oncidium ascendens Lindl. A- Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral view. F - Pollinarium and operculum. lllustration voucher: F. Pupulin 322. !OJO REVISTA DE BIOLOGIA TROPICAL becarning erect when mature, to 20 cm long, to 1 m long, many-flowered. Flowers yellow, many-flowered. Flowers small, with sepals with sepals and petals striped and blotched and petals yellow blotched with reddish-brown with reddish-brown; the lip with a large brown and the lip yellow, spotted with brown on the blotch at the base of the midlobe. Dorsal sepal callus. Dorsal sepal elliptic-oblong, obtuse, short unguiculate, elliptic, acuminate, with deeply concave, 8 mm long, 4 mm wide. undulate margins, 13 mm long, 5 mm wide. Lateral sepals obliquely oblong, lhe apex . Lateral sepals obliquely lanceolate, acute, 17 obtuse, minutely apiculate, 9 mm long, 4 mm mm long, 4.3 mm wide. Petals elliptic­ wide. Petals elliptic-obovate, obtuse, obovate, acute to acuminate, 13 mm long, 4 apiculate, 9 mm long, 6 mm wide. Lip 3- mm wide. Lip 3-lobed, widely pandurate; the lobed; the lateral lobes subquadrate elliptic, lateral lobes rounded; the midlobe without a truncate, erect, with two thickened calli at the distinct isthmus, the lamina renifonn, retuse to base; midlobe flabellate, with a distinct emarginate at apex; disc with a callus isthmus, the lamina deeply emarginate to composed by two diverging ears at the base bilobed at apex; disc with a complex callus and a central keel surrounded by two acicular fonned by a central, rounded keel surrounded projections at the middle and two apical teeth; by many short to long, acicular projections; entire lip 15 mm long, 17 mm wide across the entire lip 1.3 cm long, 1.3 cm wide between midlobe. Column slender, provided with two the lateral lobes. Column short, stout, narrowly triangular, crenulate wings, 4.5 mm provided with two elongate, incurved, long. Pollinia 2, pyrifonn, sulcate, on a narrow stigmatic arms, 3.5 mm long. Pollinia 2, stípe; viscidium elliptic, brown. Anther cap pyrifonn, sulcate, on a narrow, rhombic stipe; cucullate. viscidium elliptic, · brown. Anther cap Distribution: Epiphytíc in lowland forests obovate-oblong, cucullate, 2.6 mm long. from Mexico to Costa Rica. Distribution: Epiphytic in lowland, Parque Nacional Manuel Antonio: A disturbed forests from Mexico to Costa Rica, single collection was done not far from the and the West Indies. northern borders of the park, where the plant Parque Nacional Manuel Antonio: A was found growing on young trees in single plant was found growing on the trunk of secondary disturbed forest. This taxon is part a short tree in secondary, very disturbed forest. of a complex of closely allied species with The species is rare at PNMA. very similar morphology, including O. baueri, Phenology: flowering occurs in April­ O. isthmi, O. panamense, O. sphacelatum, and May. O. stenotis. I accept here this species as O. polycladium on the basis of the lack of a 2. Oncidium polycladium Rchb.f. ex distinct isthmus on the lip. The species should Lindl., Fol. Orch. Oncidium, 47. 1855. be considered rare at PNMA. Piant epiphytic, to 70 cm tall, with short Phenology: flowering mainly occurs in and stout rhizome. Roots fleshy, glabrous. April. Pseudobulbs clustered, ovate-elliptic, lateral! y compressed, sometimes slightly curved, 3. Oncidium stenobulbon Krzl. in strongly grooved during dry season, to 13 cm Engler, Pflanzenr. 4. 50, Heft. 80: 281. 1922. long, 4 cm wide, diphyllous, subtended by (Fíg. 30). large, scarious sheaths, with leafy limb. Phmt epiphytic, cespitose, to 20 cm tall. Leaves subcoriaceous, erect, linear, acute, Roots fleshy, glahrous. Pseudobulbs canaliculate toward the base, to 35 cm long, clustered, ovate-elliptic, laterally strongly 2.5 cm wide. Inflorescence lateral, erect, flattened, smooth, sometimes slightly curved, emerging from the axil of the leafy sheaths, with sharp edges, to 10 cm long, 7 cm wide, racemose to paniculate with short branchlets, diphyllous, subtended by large, foliaceous PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1011

5mm 5mm 1 cm

Fig. 30. Oncidium stenobulbon Krzl. A - Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral view. F- Pollinarium and operculum. Illustration voucher: F. Pupulin 303. 1012 REVISTA DE BIOLOGIA TROPICAL sheaths. Leaves thin, erect, elliptic to narrowly sepals; lip simple or 3-lobed. Column with a foot elliptic, acute, canaliculate toward the base, to at the base. Pollinia 2. 10 cm long, 4 cm wide. Inflorescence lateral, A neotropical genus of about 1,500 erect, emerging from the axil of the leafy species. sheaths, racemose to paniculate with short branchlets, to 40 m long, many-flowered. l. Pleurothallis corniculata (Sw.) Lindl., Flowers yellow, with sepals and petals striped Bot. Reg. 28: Mise. 83, 1842. Epidendrum and blotched with reddish-brown; the lip with corniculatum Sw., Prodr. 123, 1788. a large brown blotch on the isthmus. Dorsal Pleurothallis nubigena Lindl. Ann. & Mag. sepal unguiculate, elliptic, acute, with Nat. Hist. 3: 326, 1858. Pleurothallis undulate margíns, 15 mm long, 4.5 mm wide. jocolensis Ames, Sched. Orch. 2: 19, 1923. Lateral sepals shortly unguiculate, obliquely (Fig. 31). lanceolate to lígulate, acute, 18 mm long, 4 Plant small, cespitose, to 6 cm tall, with mm wide. Petals ovate to lanceolate, acute, 13 abbreviated rhizome. Roots filiform, glabrous. mm long, 6 mm wide. Lip 3-lobed, with short Ramicanls abbreviated, to about 1.1 cm long, lateral lobes, rounded to subquadrate at apex; monophyllous, covered by 3 tubular, scarious the isthmus triangular, abruptly constricted sheaths. Leaf obovate to oblong-elliptic, toward the insertion of the midlobe; midlobe coriaceous, with a short, sulcate petiole, obcordate, emarginate, with a short apicule; emarginate and obscurely tridenticulate at disc with a callus composed by a central keel apex, to 4.6 cm long, 0.8-1 cm wide. surrounded by 4 basal projectións and two Inflorescence erect, filiform, emerging from apical teeth; en tire lip 17 mm long, 11 mm wide the insertion of the leaf, 1-flowe.red, subequal across the midlobe. Column stout, provided to the leaf, to 5.5 cm long, provided with a with two obscure wing-like keels, 4 mm long. · small, scarious, acuminate bract toward the Pollinia 2, suborbicular, sulcate, on a narrow base, 2.5 mm long. Flowers small, orange. stipe with a dorsal projection; viscidium Dorsal sepal oblong-Janceolate, narrowly elliptic, brown. Anther cap cucullate. obtuse, verruculose at apex, 7.2 mm long, 2.5 Distribution: Endemic to Costa Rican mm ·wide. Lateral sepals connate almost to lowlands. the apex to form a obovate-elliptic lamina, Parque Nacional Manuel Antonio: Plants obtuse and emarginate at apex, 6.3 mm long, of Oncidium stenobulbon have not yet been 3.2 mm wide. Petals obliquely oblong, falcate observed within PNMA area. The single at the base, acute, glandulose, thickened at collection is from the low hills north of the apex, 2.5 mm long, .1.1 mm wide at the middle. park, where the plants were found on the main Lip linear-elliptic, truncate, with two small trunks of large trees in remnant of primary retrorse lobules at the base, provided at apex forest. The species should be considered rare at with two clumps of short hairs along the PNMA. margins, the disc with a central groove at the Phenology: flowering mainly occurs in base, laterally finely papillose, 2.6 mm long, January to March. 0.8 mm wide. Column with a conspicuous foot at the base, linear, winged, tridenticulate at apex, 1.8 mm long. Pollinia 2, narrowly 18. PLEUROTHALLIS R. Br. obtriangular, laterally flattened. Anther cap subtriangular, cucullate, 2-celled, 0.4 mm Cespitose or creeping herbs with long. monophyllous ramicauls at least partly Distribution: Epiphytic in dense tropical concealed by unadorned sheaths. Inflorescence forests in West Indies, Mexico, Guatemala, a terminal raceme. Sepals subequal, free or Honduras, Nicaragua, Costa Rica, and Guyana variously connate; petals generally smaller than (?). PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1013

E

1mm 3 cm Fe¡

1 mm

5 mm

3 mm 5 mm

Fig. 31. Pleurothallis corniculdta (Sw.) Lindl. A- Habit. B - Flower. C- Dissected perianth. D- Colurnn and lip, lateral view. E- Lip. F- Pollinarium and operculum. Illustration voucher: F. Pupulin 295. 1014 REVISTA DE BIOLOGIA TROPICAL

Parque Nacional Manuel Antonio: Plants 19. POLYSTACHYA Hook. were found as epiphyte on western or northwestem faces of large trunks of Brosimum Cespitose, epiphytic, rarely terrestrial utile trees in primary forest and mature secondary herbs, usually provided with definite forest. The species is frequent at PNMA. pseudobulbs. Leaves l-2, conduplicate, Phenology: flowering begins m coriaceous, persistent. Inflorescence a raceme, September through December and January. often paniculate, from the apex of pseudobulbs, many-flowered. Flowers small, 2. Pleurothallis lewisae Ames, Proc. Biol. non-resupinate. Lateral sepals larger than Soc. Wash. 44: 41, 1931. (Fig. 32). dorsal sepal, variously connate to form a Plant minute, to 7 cm tall, with creeping mentum; petals smaller than sepals, spreading; rhizome. Roots filiforn1, glabrous. Ramicauls very lip shortly clawed, en tire or 3-lobed, smooth or abbreviated, 1-3 mm long, monophyllous, covered with a conspicuous callus, often covered with by 2 tubular sheaths. Leaf obovate to broadly farinaceous cells. Column erect, semiterete, elliptic, coriaceous, abaxially finely rugulose, with a prominent foot and terminal anther. emarginate at apex, to 0.5-1.1 mm long, 3.9-7 mm Pollinia 4, waxy, sometimes joined into 2. wide. Intlorescence erect, emerging from the base A pantropical genus of sorne 200 specíes, of the leaf, consecutively 1- to 2-flowered, to 1.2 mainly African and Asían in distribution. mm long, with a small, triangular-lanceolate, acuminate bract, 0.8 mm long. Flowers small, l. Polystachya joliosa (Lindl.) Rchb.f., reddish-brown. Dorsal sepal oblong, blunty acute, Ann. Bot. Syst. t. 6: 640. 1863. Stelis foliosa 5-6 mm long, 2.5-3 mm wide. Lateral sepals Lindl., Ann. Mag. Nat. Híst. 2: 330, t. 17. slíghtly connate at the base, lanceolate-oblong, 1839. Polystachya cerea Lindl., Edward's Bot. acute, 5-6 mm long, 1.8-2.5 mm wide. Petals Reg. 26: Mise. p. 86. 1840. Polystachya oblong, obscurely apiculate, glandulose, with claváta Lindl., Edward's Bot. Reg. 20: Mise. minutely dentate margins, 4-5 mm long, 2.5 mm p. 61. 1842. Polystachya weigeltii Rchb. f. wide. Lip oblong-elliptic, with two triangular lobes Linnaea 25: 230. 1852. Polystachya below the middle, minutely glandulose, the disc caracasana Rchb. f., Bonplandia 2: 15. 1854. slightly tickened and verrucose, 2.5 mm long, 1.1 Dendrorchis caracasana (Rchb.f.) O. Ktze., mm wide between the lobes. Column with a short Rev. Gen. Pl. 659. 1891. Polystachya minar foot at the base, arcuate, winged, irregular! y den tate Fawc. & Rendle, J. Bot. 48: 106. 1910. at apex, 2.5 mm long. Pollinia 2, narrowly Polystachya altilamellata Schltr., obtriangular, laterally flattened. Anther cap Repert. Spec. Nov. Regni Veg. 10: 385. 1912. hemiglobose, cucullate, 0.4 mm long. Polystachya ecuadorensis Schltr., Repert. Distribution: A rather inconspicuous Spec. Nov. Regni Veg. Beih. 8:90. 1921. Central American species, ranging from Polystachya poeppigii Schltr., Repert. Spec. Guatemala to Panama in lowland rain forests. Nov. Regni Veg. Beih. 9: 155. 1921. Parque Nacional Manuel Antonio: Plants Polystachya guatemaltensis Schltr., Repert. were found as epiphyte, closely appressed to the Spec. Nov. Regni Veg. Beih. 17: 141. 1921. bark of large branches and limbs of Cynometra Polystachya panamensis Schltr., Repert. Spec. hemitomophylla (guapínol negro) trees in Nov. Regni Veg. Beih. 17: 49. 1922. primary forest, in very shaded conditions. The Polystachya costaricensis Schltr., Repert. species has been found only within a single plot Spec. Nov. Regni Veg. Beih. 19: 223. 1923. and should therefore be considered rare at Polystachya cubensis Schltr., Urb. Sym. Ant. PNMA. However, it is probable his very small 9: 59. 1923. Polystachya powellii Ames, size renders it easily overlooked. Sched. Orch. 7: 31. 1924. Polystachya Phenology: flowering occurs from August huebneri Schltr., Beih. Bot. Centralbl. 43, 2: through December. 112. 1925. (Fig. 33). PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1015

E

1 mm

5 mm e

~ 1mm

3 mm 1 cm

Fig. 32. Pleurothallis lewisae Ames. A - Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Pollinarium and operculum. Illustration voucher: F. Pupulin 360. 1016 REVISTA DE BIOLOGIA TROPICAL

L-·-···-·--··.::.2_m.:..;.m.;.;_ ___¡

5 mm

. e?. .

S mm 3 mm

Fig. 33. Polystachyajoliosa (Lindl.) Rchb.f. A- Habit. B- Flower. C- Dissected perianth. D- Column and Iip, lateral view. E - Pollinarium and operculum. Illustration voucher: F. Pupulin 468. PUPUL!N: Orchíd Horula of Parque Nacional Manuel Antonio 1017

Plant epiphytic, cespitose, erect, to about persistent. Inflorescence a terminal fascicle or 20 cm tall, with short, pseudobulbous stems. a raceme, 1- to few-flowered. Flowers usually Roots fleshy, glabrous. Pseudobulbs ovoid to small. Sepals and petals subequal, spreading; fusiform, enclosed in basal, papyraceous lip articulated with the column foot, straight to sheaths when young, to 3 cm long, 1.6-2 cm geniculate, with or without a claw, entire or 3- wide, bearing 3-5 leaves. Leaves oblong­ lobed. Column short, usually with a short to elliptic, to oblanceolate, subcoriaceous, prominent foot, provided with wings or articulate to the sheaths envolving the stem at auricles or wingless, the anther terminal, the base, to 20 cm long, 1.3-3 cm wide. incumbent. Pollinia 4 or 6, lateraaly Inflorescence terminal, a single, many­ compressed, ceraceous. flowered raceme or usually with severa) short A neotropical genus of sorne 50 species, branches from the nodes of the peduncle; distributed from Mexico to Brazil and Bolivia, peduncle covered with long, acuminate, and the West Indies. imbricating scarious bracts; the rhachis fuzzy­ pubescent. Ovaries sessile, finely pubescent. l. Scaphyglottis micrantha (Lindl.) Flowers small, with yellowish to pale orange Ames & Corren, Bot. Mus. Leafl. Harv. Univ. sepals and petals, the lip with a white callus 10: 85, 1942. Hexadesmia micrantha Lindl., and whitish verrucules. Dorsal sepal ovate, Bot. Reg. 30: Mise. 2, 1844. (Fig. 34). concave, acute, dorsally sparsely pubescent, Plant epiphytic, cespitose, ascending, to 3.5 mm long, 2 mm wide. Lateral sepals about 16 cm tall, with abbreviated rhizome. obliquely broadly ovate, acute, pubescent, 4 Roots filiform, slender, glabrous, with green mm long, 3 mm wide. Petals falcate-oblong, vegetative apex. Pseudobulbs fusiform, obtuse to acute at apex, 2.5 mm long, 1 mm stipitate, somewhat curved, unbranched, wide. Lip 3-lobed, oblong-ovate; laterallobes diphyllous, subtended by many scarious, small, rounded, acute at apex; the midlobe imbricating sheaths, to 7 cm long, 0.8 cm wide. suborbícular, truncate, retuse, farinose on the Leaves linear, obliquely retuse, to 8.4 cm long, inner surface; disc provided with a mamillate 0.8 cm wide. Inflorescence terminal, callus at the base; entire lip 3.5 mm long, 3 mm racemose, many-flowered, to 8.5 cm long; wide. Column stout, with a short foot, about 1 peduncle filiform, provided with severa!, mm long. Pollinia 4, on a short caudicle. spreading sheaths, subtended by severa! Anther cap cucullate, somehath triangular, 2- scarious sheaths. Ovaries pedicellate, linear, celled. abruptly thickened at apex. Flowers very Distribution: A widespread species in the small, with white sepals and petals, and green Neotropic, distributed from Mexico to South lip and column. Dorsal sepal elliptic-obovate, America and the West Indies. obtuse to slightly apiculate, concave, 2 mm Parque Nacional Manuel Antonio: Plants of long, 1 mm wide. Lateral sepals obliquely Polystachya foliosa are locally common in broadly ovate-elliptic, apiculate, 1.9 mm long, secondary vegetation along the eastem border of 1.4 mm wide. Petals obliquely elliptic, rounded the park. However, it has not yet been reported and apiculate at apex, 1.7 mm long, 0.9 mm from PNMA, where it is regarded as rare. wide. Lip 3-lobed, subreniform-flabellate, 1.6 Phenology: flowering occurs m mm long, 2.5 mm wide; the lateral lobes September and October. obliquely elliptic, broadly rounded at apex, erect and surrounding the column in natural 20. SCAPHYGLOTTIS Poepp. & Endl. position; the midlobe transversally elliptic to Cespitose or repent, epiphytic herbs, with subquadrate, apiculate, with minutely crenulate definite pseudobulbs or thickened, simply or margins. Column short, ten~te, slightly arcuate, fasciculated, branched stems. Leaves 1-3, 1.2 mm long. Pollinia 6, on a short caudicle. conduplicate to semiterete, thin to coriaceous, Anther cap subcordate, cucullate, 2-celled. 1018 REVISTA DE BIOLOGIA TROPICAL

2 mm

2. mm

5 cm

E

3 mm 1mm

Fig. 34. Scaphyglottis micrantha (Lindl.) Ames & Correll. A- Habít. B - Flower. C - Díssected perianth. D - Column and lip, lateral view. E - Operculum and pollinaria. Illustration voucher: F. Pupulin 377. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1019

Distribution: A Central American species mm long, 3.5 mm wide. Column terete, distributed from Guatemala to Panama. provided at the base with a hole to enter the Widespread in Costa .Rica. pedicellate spur, about 3 mm long. PoHinia 4, Parque Nacional Manuel Antonio: Plants on two short, bifid caudicles. Anther cap were found as epiphyte on Hymenaea rounded, cucullate, 4-celled. courbaril (guapinol) in disturbed primary Distribution: A widespread species forest. The species is occasional at PNMA. distributed from lowlands to about 1,000 m Phenology: flowering recorded in July. from Guatemala to Bolivia and Peru. Parque Nacional Manuel Antonio: Plants 2. Scaphyglottis prolifera Cogn., Mart. were found as common epiphyte on Citrus spp. Fl. Bras. 3(5): 15. 1898. Scaphyglottis cuneata in orchards close to the park. It has not yet Schltr., Beih. Bot. Centralbl. 36: 398. 1918. been reported from PNMA, where it should be Tetragamestus gracilis Schltr., Beih. Bot. regarded as a rare species. Centralbl. 36: 400. 1918. Scaphyglottis Phenology: flowering occurs from gracilis (Schltr.) Schltr., Fedde Rep. Sp. Nov. November to February. Beih. 19: 28. 1923. Scaphyglottis werklei Schltr., Fedde Rep. Sp. Nov. Beih. 19: 28. 3. Scaphyglottis stellata Lindi., Bot. Reg. 1923. Ponera mapiriensis Krzl., Fedde Rep. n.s. 12: mise. 60. 1839. Ponera amethystina Sp. Nov. Beih. 25: 22. 1928. (Fig. 35). Rchb.f., Saunders Ref. Bot. 2: t. 93. 1869. Scaphyglottis amethystina (Rchb.f.) Schltr., Plant epiphytic, cespitose, ascending, to Beih. Bot. Centralbl. 36 (2): 456. 1918. about 20 cm tall, with pseudobulbous, usually Scaphyglottis brachiata Schltr., Fedde Rep. superimposed stems. Roots filiform, slender, Sp. Nov. Beih. 9:432. 1911. (Fig. 36). glabrous, emerging from the rhizome or from the insertion of new growths at the apex of old Plant epiphytic, cespitose, erect to pseudobulbs. Pseudobulbs linear, terete, pendent, to about 40 cm tall, with sometimes slightly attenuate at both the apices, pseudobulbous, usually superimposed to 10 cm long, 0.4 cm wide, diphyllous; two segments. Roots filiform, slender, glabrous, new, shorter pseudobulb~ usually develop the emerging from the rhizome or at the connection next year at the apex of old pseudobulbs, of old and new pseudobulbs. Pseudobulbs emerging from short, imbricating, papyraceos fusiform, stipitate, to 12 cm long, 0.5 cm wide, sheaths. Leaves linear-elliptic, subcoriaceous, diphyllous, each pseudobulb usually bearing a obliquely retuse, to 4.5 cm long, 0.5 cm wide. single new pseudobulb at apex the next year, Inflorescence terminal at the apex of each emerging from imbricating, papyraceos internode of the stem, a single to few-flowered sheaths. Leaves linear-lanceolate to narrowly fascicle; peduncle subtended by severa!, elliptic, obliquely retuse, to 12 cm long, 0.8 cm imbricating, scarious sheaths. Ovaries wide. Inflorescence a few-flowered fascicle pedicellate, linear-clavate. Flowers very small, born at the apex of each internode of the stem. with greenish sepals and petals, the lip white Ovaries pedicellate, linear. Flowers small, striped with purple and purple-brown anther. with white sepals and petals, the lip rose-purple Dorsal sepal linear-elliptic, acute, slightly striped with purple; column and anther deep concave toward the apex, to 5 mm long, 2.5 purple. Dorsal sepal lanceolate-elliptic, acute, mm wide. Lateral sepals obliquely elliptic, slightly concave toward the apex, 7 mm long, acute to obtuse, 6 mm long, 3 mm wide. Petals 2.5 mm wide. Lateral sepals connate at the ligulate, rounded and apiculate at apex, 4 mm base, forming a short mentum, obliquely linear­ long, 1 mm wide. Lip spathulate-flabellate, lanceolate, acute, 8 mm long, 2.5 mm wide. emarginate, with a short apicule, slightly Petals ligulate, slightly constricted toward the concave toward the base, reflexed at apex, 6 apex, then rounded and apiculate, 7 mm long, 1020 REVISTA DE BIOLOGIA TROPICAL

5 mm

F

zmm

5 cm

3 mm

5 mm 3 mm

Fig. 35. Scaphyglottis prolifera Cogn. A- Habit. B - Flower. C- Dissected perianth. D- Coturno and lip, lateral view. E­ Column, ventral view. F - Pollinaria and operculum. Illustration voucher: F. Pupulin 471. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1021

1 cm

10 cm

5 mm 2mm 3 mm

Fig. 36. Scaphyglottis stellata Lindl. A - Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral view. F- Pollinaria and operculum .. lllustration voucher: F. Pupulin 300. 1022 REVISTA DE BIOLOGIA TROPICAL

1.5 mm wide. Lip clawed, the blade 3-lobed, covered with amplexicaul leaf-sheaths, obovate-flabellate; laterallobes large, rounded, sometimes branched, lepidote to slightly erose at apex, inserted at the middle of the lip; furfurescent. Roots fleshy, glabre. Leaves midlobe subquadrate, obtuse to emarginate; plicate-veined, lanceolate to lanceolate-ovate, entire lip 7 mm long, 5 mm wide between the acuminate, coriaceous, to about 15 cm long, 3- lateral lobes. Column terete, slightly arcuate, 5 cm wide. Inflorescence apical, sessile, a 1- wíth two riarrowly triangular stelidiar arms flowered raceme. Flowers large, not toward the apex, 3 mm long. Poninia 4, on completely spreading, rose-purple with darker short bifid caudícles. Anther cap cucullate, lip. Sepals free, more or less reflexed at the tip. rounded, 4-celled. Dorsal sepal lanceolate, acute, slightly Distribution: A common species in concave, about 6 cm long, 1.6 cm wide. lowlands in Costa Rica and Panama to Guianas Lateral sepals obliquely lanceolate, acute, and Bolivia. concave toward the apex, 6 cm long, 1.4 cm Parque Nacional Manuel Antonio: Plants wide. Petals elliptic-lanceolate, acute, 5.7 cm of Scaphyglottis stellata are common long, 2.2 cm wide. Lip obovate, obtuse to epiphytes in al! types of vegetation at the Park, minutely retuse, deeply concave, the basal where they often form big clumps of many margins forming a tu be enclasping the column, stems, usually pendent in mature specimens. It the apex deflexed, with slightly crenulate is a very common species at PNMA. margins, 6.5 cm long, 3.6 cm wide; callus at Phenology: flowering occurs from the base of the lip formed by two low, slender, October to January. divergent carines. Column elongated, subterete, slender, abaxially keeled, to 3.5 cm long, with two introrse teeth at apex. Pollinia 21. SOBRALIA Poepp. & Endl. 8, subceraceous. Anther cap cucullate, bifid at the base, about 3 mm long. Terrestrial or epiphytíc, small or large Distribution: A relatively common species herbs, with foliaceous, reed-like stem. Leaves in lowlands from Mexico to Panama. sessile, pergameneous, plicate. Inflorescence a Parque Nacional Manuel Antonio: terminal raceme, often 1-flowered. Flowers Stobralia decora is known from a few plots at usually large, fugaceous. Sepals and petals PNMA, where it is usually restricted as subequal, sometimes shortly connivent at the epiphyte to old trunks and large branches in base, spreading, the petals usually broader than primary or disturbed primary forest. The sepals; lip not clawed, entire or 3-lobed, species is occasional at PNMA. usually concave, the basal margins enfolding Phenology: flowering recorded m the column, the disc smooth or lamellate, December. usually with a inconspicuous to conspicuous ca!lus at the base. Column elongate, usually ventrally provided with a strong keel, footless, 22. STENORRHYNCHOS L.C. Rich. ex anther incumbent. Pollinia 8, granular or Spreng. subceraceous. A neotropical genus of about 100 species, Ten-estría! or epiphytic herbs with or widespread from Mexico to South America, without leaves at flowcring, with basalleaves. and the West Indies. Leaves short stalked, lanceolate, deciduous. Inflorescence erect, a usually dense raceme l. Sobralia decora Batem., Orch. Mex. & with colored bracts, many-flowered. Flowers Guat. t. 26. 1841. (Fig. 37). relatively large. Sepals and petals subequal, the Plant epiphytic or rarely terrestrial, erect connivent bases of sepals forming a tube, to pendent, to about 2 m tall. Stem slender, spreading at apex; lip not clawed, the margins PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1023

scm

5 cm

;: .: .• ~- F

3cm 1 '\cm 5mm

Fig. 37. Sobralia decora Batem. A- Habit. B - Flower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral and lateral view. F - Operculum. Illustration voucher: F. Pupulin 320. 1024 REVISTA DE BIOLOGIA TROPICAL thickened at the base, with or without a long. PoUinia 2, narrowly pyriform, on a constriction in the middle. Column massive, ligulate viscidium enveloping the rostellum. provided with a long foot, anther dorsal. Anther cap acutely triangular, persistent, Pollinia 2, powdery. about 3 mm long. A small neotropical genus of sorne 3 Distribution: A widespread species species, widespread from Florida to Argentina, distributed throughout tropical America. and the West Indies. Parque Nacional Manuel Antonio: Stenorrhynchos lanceolatum is known from a l. Steno"hynchos lanceolatum (Aubl.) single collection at Jardín Gaia, not more than L.C. Rich. ex Spreng., Syst Veg. 3: 710. 1826. 3 km west of the park along the road to Limodorum lanceolatum Aubl., PI. Guian. 2: Quepos. Here a small population was found in 821. 1775. Satyrium orchioides Sw. Prodr. Veg. rocky soil in secondary vegetation. :rhe species Ind. Occ. 118. 1788. Neottia lanceolata (Aubl.) should be regarded as rare at PNMA. Willd., Sp. PI. 4: 75. 1805. Spiranthes orchioides Phenology: flowering recorded m (Sw.)A. Rich. in Sagra, Hist. Fis. Poi. Nat. Cuba November to December. pt. 2, 11 [Bot.]: 252. 1850. Gyrostachys lanceolata (Aubl.) O. Ktze., Rev. Gen. PI. 2: 664. 1891. Spiranthes jaliscana S. Wats., Proc. 23. TRIGONIDIUM Lindl. Amer. Acad. 26: 153. 1891. Spiranthes lanceo/ata (Aubl.) Leon, Contr. Ocas. Mus. Hist. Cespitose, epiphytic, erect to scandent Nat. Col. de la Salle, Acta 8: 385 .. 1946. Sacoila herbs with sulcate pseudobulbs, 2- to 5-foliate lanceo/ata (Aubl.) Garay, Leafl. Bot. Mus. at apex. Leaves conduplicate, linear, Harvard Univ. 28: 352. 1982. (Fig. 38). subcoriaceous, persistent. Inflorescence a simple, short to usually elongate raceme from Plant terrestrial, erect, to about 30 cm tall, the bases of pseudobulbs, 1-flowered. Flowers with basal, soft, herbaceous leaves, 1eafless at relatively large. Sepals subequal, the bases flowering stage, the stem enveloped by severa} connivent forming a tube, the apex spreading; .amplexicaul bracts. Roots fleshy, clustered. petals smaller than sepals, with an ·apical, Leaves 3-6, in a basal rosette, glossy callus; lip 3-lobed, the lateral lobes membranaceous, elliptic, 15-19 cm long, 2-3.5 erect, the midlobe thickened, the disc with a cm wide. Inflorescence a congested, ligular callus. Column short, semiterete, pubescent, many-flowered raceme to 50 cm footless, with terminal, incumbent anther. long, with attenuate bracts. Flowers small, not Pollinia 4, waxy. completely spreading, pubescent on the A small neo tropical genus of sorne 1O eitemal surface, rose-orange with white to species, ranging from Mexico to Brá.zil. pale rose lip. Sepals more or less connivent, spreading at the tip. Dorsal sepal lanceolate, l. Trigonidium egertonianum Batem. ex acute, dorsally pubescent, about 20 mm long, 6 Lindl., Bot. Reg. n.s. 1: Mise. 73. 1838. mm wide. Lateral sepals lanceolate, acute, Trigonidium seemanni Rchb.f., Seem. Bot. dorsally pubescent, the base decurrent on the Voy. Herald 214. 1854. (Fig. 39). ovary producing an obtuse mentum, 23 mm Plant epiphytic, cespitose, forming large long, 4.5 mm wide. Petals linear-elliptic to clumps. Roots fleshy, glabrous. PseudobUlbs Ianceolate, falcate, acute, 18 mm long, 4 mm ovate, somewhat laterally compressed, sulcate, wide, coherent to the dorsal sepal. Lip to 3.5 cm long, about 3.2 cm wide, basally lanceolate, acute to obtuse, the apex deflexed, covered by 2-3 papyraceous sheaths, bifoliate 23 mm long, 7 mm wide, the margins erect, at apex (rarely monophyllous). Leaves fleshy, enclosing the column. Column wide narrowly oblanceolate to linear, arching, acute, basally, with an acuminate rostellum, to 6 mm to about 30 cm long, 1.3-2 cm wide. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1025

1 cm

3 cm

'------3cm

F

5mm

1 cm 10 cm L---.----··-----'

Fig. 38. Stenorrhynchus lanceolatum (Aubl.) L. C. Rich. ex Spreng. A- Habit. B- Inflorescence. C- Flower. D • Dissected perianth. E- Column and lip, lateral view. F- Column, lateral view (pollinarium removed). G- Operculum and pollinarium. lllustration voucher: F. Pupulin 356. 1026 REVISTA DE BIOLOGIA TROPICAL

5mm

3 mm

o

1 crn 3 cm

Fig. 39. Trigonidium egerErmianum Batem. ex Lindl. A- Habit. B - F1ower. C - Dissected perianth. D - Column and lip, lateral view. E - Column, ventral view. F - Operculum and pollinarium. Illustration voucher: F. Pupu/in 368. PUPULIN: Orchid Florula ofParque Nacional Manuel Antonio 1027

Inflorescences basal, erect, slender, single­ deeply concave, the laterals connate nearly to flowered scapes to 20 cm long, covered by the apex, forming a mentum at the base; petals many closely appressed, tubular, papyraceous about as long as sepals; lip shortly clawed, 3- sheaths. Flowers rather showy, pale yellow to lobed, articulated with the column base, the pale orange striped and reticulated with purple, apex fleshy. Column erect, subcylindric­ the petals provided with metallic blue to clavate, with terminal, incumbent anther. mauve blotch near the apex. Sepals free, Pollinia 2, waxy. appressed toward the base forming a tube, then Probably .a monotypic neotropical genus, abruptly expanded and reflexed at apex. widespread from Costa Rica to Trinidad, Dorsal sepal shortly unguiculate, elliptic" Brazil, Bolivia and Peru. oblanceolate, acute, 3.3 cm long, 1.8 cm wide. Lateral sepals obliquely ellíptic-lanceolate, l. Trizeuxis falcata Lindl., Collect. Bot. t. acute, 3.4 cm long, 1.3 cm wide. Petals 2. 1823. Trizeuxis andina Schltr., Repert. Sp. lanceolate, acute, about 1.8 cm long, 0.6 cm Nov. 10: 52. 1922. (Fig. 40). wide. Lip 3-lobed, articulate with the base of Plant epiphytic, small, erect, to about 5 the column; lateral lobes erect and parallel to cm tall, with a short, distichously foliaceous the column, acute; the midlobe ovate, acute to stem. Roots flexuous, glabre. Pseudobulbs obtuse, rather fleshy, verrucose; disc with a rounded to subquadrate, to 5 mm long, about 4 fleshy ligulate callus as long as the lateral mm wide, basally covered by 2-3 imbricating lobes; entire lip 0.8 cm long, 5 mm wide leafy sheaths, monophyllous. Leaf fleshy, between lateral lobes. Column subterete, laterally flattened, falcate to gladiate, about 7 mm long. Pollinia 2, elliptic, on a acuminate, 1-3.5 cm long, 0.2-0.4 cm wide, saddle shaped caudicle. Anther cap cucullate, articulate to the basal sheaths and arranged like keeled. a fan. Inflorescence a basal, arcuate to Distribution: A widespread species from pendent, slender, paniculate raceme to 4 cm Mexico to Panama and Colombia, very long, the lateral subcapitate branches about 2 common in Central American lowlands. cm long, many-flowered. Flowers very small, Parque Nacional Manuel Antonio: The subglobose, greenish yellow to pale yellow species is usually restricted to primary and with yellow to orange lip. Dorsal sepal ovate, secondary mature forest, where it establishes retuse to emarginate, deeply concave, partially itself in partial shade on trunks and large limbs connate with the petals, 1.5 mm long, 1.2 mm covered with bryophytes. Though common wide. Lateral sepals connate to nearly the through its range, the species is only apex to produce a obovate, bifid synsepalum, occasional at PNMA. 1.9 mm long, 1.5 mm wide. Petals elliptic­ Phenology: flowering occurs from ovate, obtuse, about 2.5 mm long, 1.2 mm November to February. wide. Lip 3-lobed, lanceolate, acute, concave at the base, with erect lateral margins enclosing the column, the apex fleshy, about 24. TRIZEUXIS Lindl. 2.5 mm long. Column short, stout, subterete, dilated at apex, to 1 mm long. Pollinia 2, Cespitose, epiphytic herbs with short, pyriform, on a large stipe. Anther cap distichously foliaceous stem, the leaves cucullate, very large relatively to the column, arranged in a fan envolving a small about 1 mm long. pseudobulb, 1-foliate at apex. Leaves falcate, Distribution: A widespread species in laterally flattened, coriaceous. Inflorescence a South America to Brazil and Bolivia., reaching pendent, paniculate raceme from the base of in Costa Rica its northem distributionallimit. pseudoblb, many-flowered. Flowers Parque Nacional Manuel Antonio: The subglobose, small. Sepals dissimilar, the dorsal species is known only from bushes and 1028 REVISTA DE BIOLOGIA TROPICAL

5mm L ...... ············-····----········-__J

F

1 mm

3mm

Fig. 40. Trizeuxis fa/cata Lindl. A - Habit. B - Aower. C - Apex of inflorescence. D - Dissected perianth. E - Column and lip, lateral view. F - Pollínarium and operculum. Illustration voucher: F. Pupulin 473. PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1029 orchards outside the park, about lkm westthe a raceme to about 15 cm long, many­ border of PNMA along the road to Naranjito. flowered; bracts lafice6}ate to ovate, obtuse, Though usually common through its range, cucullate, l cm long. Flowers large and Trizeuxis falcata should be regarded as rare at fleshy, greenish yellow to pale yellow with PNMA. white lip. Dorsal sepal linear elliptic to Phenology: flowering most of the year. In oblanceolate, acute to obtuse, concave, 7.5 Ecuador the species is pollinated by small bees cm long, 1.3 cm wide. ~lateral sepals of the genus Trigona (Dodson & Marmol­ obliquely and narrowly oblanceolate, acute Dodson 1980): to obtuse, to about 8 cm long, 1.5 cm wide. Petals narrowly elliptic, obtuse to acute, about 7 cm long, 1.2 cm wide; Lip clawed, 25. VANILLA Sw. the isthmus adnate to the column, obovate to subrhombic, trumpet shaped, retuse to Epiphytic, scandent, leafy or aphyllous emarginate, concave at the base, crenulate herbs, with elongate, erect to pendent, often toward the apex, the disc with a ~uft of hairs, branched stems. Leaves conduplicate, about 8.5 cm long, 3.5-4 cm wide toward the coriaceous or .fleshy, persistent, sessile or with apex. Column subterete, slender, without a a short petiole. Inflorescence a short raceme foot, slightly dilated at apex, to 6 cm long~ from the axils of the leaves, usually with many Polleo mealy. Anther cap ventral, cucullate, flowers produced successively. Flowers about 7 mm long. Fruit a fusiform capsule usually large. Sepals and petals subequal, (the "bean") about 15 cm tongO· spreáding, free; the lateral sepals sometimes Distribution: A widespread species from shortly connate; lip clawed, simple or 3-lobed, Mexico to South America, often éultivated as the claw adnate to the column, the laterallobes ornamental and a s6urce of flavouring. erect to •enfold the colum, the midlobe Parque Nacional Manuel · Antonio: spreading or ·reflexed, usually provided with Cultivated as an ornamental at Jardín Gaia, a longitudinal keels or lines of papillae and with few kilometers from the Park ori the road to a tuft of hairs at the base. Column elongate, Quepos. Natural populations ofthis species are withou~ a foot, wingless, with incumbent said to be found in the neighbpuring remnants anther. Pollinia granular. of primary forest. The spe,cies should be The only member of the Orchidaceae regarded as rare at PNMA. having a truly commercial value, Vanilla is a Phenology: flowering occurs in March to pantropical genus of about 100 species widely April. cultivated for the flavouring extraer from its seed pods. Excluded species; The following species were reported for PNMA (Bolaños et al. l. Vanilla pompona Schiede, Linnaea 4: 1983), but no specimens ·are conserved in 573, 1829. Vanilla pompona Lindl., Gen. & Costa Rican herbaria and actual plants were Sp. Orch. PI. 437. 1840. Vanillapittieri Schltr., not found by the' authcír either within the Park Fedde Rep. Spec. Nov. 3: 106. 1906. or in the neighbouring ateas: Plant emiepiphytic, large, scandent, to Bletia purpurea (Lam.) DC. Outside the 10m tall, with a large, branched, foliaceous study area this speCies was found just to rocky stem. J,toots glabre. Leaves variable, from beachs in southern Pacific coast at Isla Vrrolíri, narrowly elliptic to lanceolate-ovate, fleshy Puntarenas (Poveda & G6mez 4717, CR!). to coriaceous, acute' or obtuse, 10-30 cm Brassia caudata (L.) Lirid[ No actual long, 3-8.5 cm wide, ·articulate to the terete specimens of this taxon were obserted from stem. Inflorescences lateral, .produced form PNMA nor in museum collections. Clairns for the axil of the leaves, erect to arcuate, stout, the existence of Brassia spp. has been reported 1030 REVISTA DE BIOLOGIA TROPICAL

(J.B. García, pers. comm.), but due to the uncata are usually small, and their presence great tall of this species and the showy may be easily overlooked. flowers, it is unikely it has escaped attention Myrmecophyla tibicinis (Batem.) Rolfe until now. (as Schomburgkia). Due to the large size of the Cyrtopodium punctatum Lindl. This plants of this species, it is unlikely that they species is generally associated to dry, strongly have escaped attention by botanists and seasonal vegetation, and its presence at PNMA aficionados. No specimens were observed at is unlikely. any of the studied plots. Chysis aurea (L.) Lindl. This species is Mormodes igneum Lindl. No collections not actually reported for Costa Rica. of this taxon from PNMA are known, but M. Encyclia cordigera (H.B.K.) Dressler (as fractijlexa Rchb.f. has been collected from Epidendrum atropurpureum). This species is southemmost Península de Osa (C. Skotak s.n., generally associated to seasonal vegetation and USJ!). ecological conditions at PNMA are unlikely to Oncidium ampliatum Lindl. Tliough this meet species requirements. species is typical of deciduous forests (Janzen, E. ionophlebia (Rchb.f.) Dressler (as 1983), an early report exists from Manuel Epidendrum ionophlebium). E. ionophlebia is Antonio area, where the species lived mixed a species usually restricted to mountainous with Laelia rubescens populations (J.B. areas above 800 meters elevation, and its García, pers. comm.). occurence at PNMA is unlikely. However, the Oncidium dichromatichum (as O. presence in our area of members of Encyclia cabagrae Schltr.). This taxon is generally subgenus Osmophytum, like E. chacaoensis, is found from about 800 to 2500 meters above highly probable. sea level, and its presence at PNMA is Laelia rubescens Lindl. Although this unlikely. lt is likely the juvenile pseudobulbs species has not been collected recently, of Aspasia epidendroides, blotched with undocumented records exist of early purple, were misinterpreted as this species. collections in the area surrounding the Park (J. Psygmorchis pusilla (L.) Dodson & B. García, pers. comm.). Dressler (as Oncidium pusillum). Though Lockhartia acuta (Lindl.) Rchb.f. Reports common on disturbed vegetation and orchards, of this species from Manuel Antonio are the presence of this species at PNMA has not probably based on the misidentification of yet been confirmed. It may be previous claims specimens pertaining to L. pandurata Pupulin. were based on observations of the similarly L. pittieri Schltr. This species is restricted psygmoid Trizeuxis fa/cata. to the Atlantic (Caribbean) lowlands, and its Stanhopea pulla Rchb.f. This species is presence at PNMA is unlikely. native from the Caribbean lowlands. Maxillaria alba (Hook.) Lindl. Due the Stelis sp. Besides Pleurothallis large size of this species, it is unlikely it have comiculata and the diminutive P. lewisae, no been so long overlooked at PNMA. other member of the Pleurothallidinae were M. planicola C. Schweinf. No specimens found within the Park area. of this taxon were observed within the studied Trichopilia maculata Rchb.f. The plots. No collections from the study area were presence of thís species at such low elevation found in Costa Rican herbaria. A very similar is very unlikely. species has recently been described (Atwood 1995) from Monteverde and the Atlantic slopes of Volcán Barva. ACKNOWLEDGMENTS M. uncata Lindl. Though no actual specímens of thís taxon were found at PNMA I am grateful to José Antonio Salazar or in the neíghbouring areas, plants of M. Alvarez, Director of PNMA, for his courtesy PUPULIN: Orchid Florula of Parque Nacional Manuel Antonio 1031 and logistic support and to Delio Salazar Chase, M.W. 1987. Obligate twig epiphytism in the Ribera, Servicio de Parques Nacionales, for his Oncidiinae and other neotropical orchids. Selbyana continuous help as a fieldguide during 10:24-30. sampling and collection of botanical material. Chase, M.W. & J.S. Pippen. 1988. Seed morphology in the 1 am very much indebted to Dario Oncidünae and related subtribes. Syst. Bot. 13: 313- Castelfranco, Director, Jardín Gaia Botanical 323. Garden, for the many facilities and for his Dodson, C.H. & P. Marmoi-Dodson, 1980. Trizeuxis suggestions and critica! comments during all falcata, Icon. PI. Trop. 4: sub pl. 350. stages of the work. Joaquín B. García shared with me bis life-long acquaintenance with Dressler, R.L. 1966. Observations on orchids and Costa Rican orchids, and Robert L. Dressler euglossine bees in Panama and Costa Rica. Rev. gave me bis valuable advice for the Biol. Trop. determination of difficult taxa. This paper is Dressler, R.L. 1993. Field guide to the orchids of dedicated to my daughters, Margherita and Costa Rica and Panama. Comstock, lthaca, New Carlotta, for whatever it cost us. York.

Holdridge, L. R. 1982. Ecología (basada en zonas de vida). REFERENCES Instituto Interamericano de Cooperación para la Agricultura, San José, Costa Rica. Allen, P.A. 1952. The swan orchids, a revision of the genus Cychnoches. Orch. J. 1: 1-226. Janzen, D.H. (ed.) 1983. Costa Rican natural history. Chicago University, Chicago. Atwood, J. T. 1995. 1\vo overlooked species of Maxillaria from Central America. Selbyana 16: 242-245. Mora-Retana, D. E. & J. B. García. 1992. Lista actualizada de las orquídeas de Costa Rica (Orchidaceae). Bolallos, R., R. Campos, M. Hammond, R. DuBois, C. Brenesia 37: 79-124. Esquive!, L. Gómez, M. Hatzüolos, L. Moreira & J. Mora. 1982. El Parque Nacional Manuel Antonio. Rojas, M., G. Robitaille, J. R. Barborak, F. Carr, R. Inventario Biologico terrestre y marino. Estudio Morales, C. McFarland & J. Calderón. 1983. Plan de oceanográfico y diseño paisajista. Centro Científico manejo y desarrollo Parque Nacional Manuel Tropical, San José, Costa Rica. Antonio. Centro Agronomico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Boza, M. A. 1986. Parques Nacionales Costa Rica Costa Rica. National Park:s. Fundación de Parques Nacionales, Costa Rica. lncafo, Madrid. Tosi, J. 1969. Mapa ecológico de Costa Rica. Centro Científico Tropical, San José, Costa Rica. Campos, J. (coordinator). 1983. Propuesta plan de manejo Parque Nacional Manuel Antonio. Resumen Wong Reyes, G. 1990. Ecologia del Mono Titi:Saimiri ejecutivo. FUNDENI, Servicio de Parques Nacio­ oerstedi cltrinellus en el Parque Nacional Manuel nales and Instituto Costarricense de Turismo, San Antonio, Costa Rica. Tesis de Grado, Universidad José, Costa Rica. Nacional, Heredia, Costa Rica.