©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at Ann Naturhist Mus Wien, B 112 239-252 Wien, März 2011 Illustrations and Studies in Neotropical Orchidaceae A note on Pleurothallis luctuosa (Pleurothallidinae), with a new species F Pupulin*’ **’ ***, D B ogarin*’ ** & M Fem ändez* A b stra c t The identity o f Pleurothallis luctuosa, its taxonomic history and geographic distribution are discussed, and the species is described and illustrated A new species from C osta Rica, P neglecta, is described and illustrated Pleurothallis neglecta is com pared w ith P luctuosa, from w hich it differs by the distinct conduplicate leaves, different color o f the flowers, triangular and smaller lip, falcate, erose petals, and the callus formed by two conspicuous, densely papillose keels Key words: Orchidaceae, Pleurothallidinae, Pleurothallis, Acronia, Pleurothallis luctuosa, Pleurothallis neglecta, Costa Rica, A.R Endres, new species Introduction In 2008, the Lankester B otanical Garden (LBG) at the U niversity o f C osta R ica began a large project devoted to the reconstruction o f the phylogenetic relationships and the assessment o f the taxonom ic identity o f the species o f subtribe Pleurothallidinae L i n d l (Orchidaceae) in C osta Rica Pleurothallidinae represents the m ost diverse orchid group in the N eotropics, and the taxonom y o f this large assem blage o f species has been traditionally problem atic, both under the profiles o f generic circum scriptions and taxa identification P leurothallis subgen Pleurothallis sect Pleurothallis subsect Acroniae (P r e s l ) L u e r (= A cronia P r e s l pro parte, L u e r 2005) includes in C osta R ica a relatively sm all n u m ber o f species W ith som e sixty described taxa, the subsect Acroniae (sensu L u e r 1998) is predom inantly South A m erican in distribution, w ith only six species recorded from C osta Rica In revising specim ens o f this group in the living collection o f LBG, we noted that the nam e P leurothallis luctuosa R c H B f w as apparently applied to two different taxa, one o f w hich is still undescribed Taxonomic history of Pleurothallis luctuosa The circum stances, under w hich A.R Endres died in Colom bia, supposedly in the first months o f 1875, have been traditionally unclear, until a recent article brought some light Franco Pupulin, Diego Bogarin, Melania Fernandez, Lankester Botanical Garden, University o f Costa Rica, P.O Box 302-7050 Cartago, Costa Rica ^ Angel A ndreetta A ndean O rchids R esearch Center, U niversity A lfredo Perez G uerrero, G ualaceo, Ecuador Harvard University Herbaria, Cambridge, MA, U.S.A - franco.pupulin@ ucr.ac.er 240 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B ] p on the crucial facts o f his biography ( O s s e n b a c h et al 2010) Endres left Costa Rica, the country where he spent the last eight years o f his life, in the course o f 1874, when he traveled to Europe and m et H einrich G ustav R eichenbach in Ham burg According to R e i c h e n b a c h (1875), they had good days talking about orchids in the Company of B enedikt R oezl (1 -1 8 ), the C zech explorer, gardener and botanist, who was considered the greatest orchid collector o f his time B y the end o f the summer, Endres crossed the A tlantic again, and in late A ugust he was in N ew York A few months later he died o f pleurisy in the C aribbean village o f Dibulla, at the foot o f the Sierra Nevada de Santa M arta m ountains in Colom bia The prestigious G ardener’s Chronicle presented the sad new o f E n d re s’ death on M ay 8th, 1875, prom ising an obituary o f the great orch id olo g ist by Prof R e i c h e n b a c h ( A n o n y m o u s 1875), but this text was never published, and likely it w as never written By the tim e the destiny was driving Endres to the date w ith his death in Dibulla, a new Pleurothallis flow ered for the first tim e in the orchid collection o f the B otanical Garden o f the U niversity o f Ham burg, o f w hich H.G R eichenbach had been appointed director in 1862 In the original description o f the new species published in Linnaea, R e ic h e n b a c h (1877) m ade no reference to Endres and the origin o f the plant, nor he gave any reasons to baptize the new Pleurothallis as “luctuosa” , or m oum ful, ju st recording that he saw the living plant in the collection o f the H am burg B otanical Garden M any years later, S c h l e c h t e r (1923) strongly protested for the lack o f inform ation associated with m any o f the new species that R eichenbach published in Linnaea H owever, in 1878, R eichenbach augm ented the description o f Pleurothallis luctuosa and illustrated it in X enia O rchidacea (Fig 1) and here he credited Endres as the discoverer o f the new species, explaining that he was ju st exam ining the first opening, black-purple flowers, w hen he was su rp rised by the new s o f the death o f his friend “H ence the name” (R e i c h e n b a c h 1878) A ccording to the m anuscript notes by Endres w hich are now in Vienna, P luctuosa had been found in C osta R ica ten years before, in 1867 In his m eticulous style, Endres prepared a detailed description and a set o f drawings o f his finding, including the habit o f the plant and floral dissections, as well as a series o f duplicate specim ens under his num ber 50 It has to be noted that this num ber has not to be interpreted as a “collecting num ber” in the m odern sense; instead, Endres m ostly used his num eration system to refer to taxa rath er than to individual collections A ctually, only one o f the sheets conserved in Vienna is labeled as P luctuosa in R eichenbach’s handw riting [W-Rchb Orch 53791!], and the m ounted specim en w as likely p repared from living material cultivated in Ham burg The holotype sheet includes several inflorescences, ramicauls and detached leaves, all o f them apparently prepared in H am burg from the living collection accessio n N° 2537, plus sketches by R eichenbach h im se lf and a pencil drawing by Endres w ith a note about the color o f the plant and the flowers, and a record o f the date and altitude o f the original collection (Endres 50) Technically, there is no w ay to uneq u iv o cally associate the living plant accessed to the H am burg Botanical Garden collection w ith E ndres’ collection bearing his num ber 50, and we consider more correct to eite the holotype o f P luctuosa as Endres sine num ero, even though the plant flowered in H am burg w as likely part o f the original collection m ade by Endres at San Ramon ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at pur■ULIN et al.: A note on Pleurothallis luctuosa (Orchidaceae) 241 m Fig 1: Drawing of Pleurothallis luctuosa, from Xenia Orchidacea, 1878 242 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B / /-> v• Fig 2: Drawing of Pleurothallis luctuosa by A R Endres [W-Rchb Orch 33417], The illustrations sent from C osta R ica [W-Rchb Orch 33417!, 36294!] (Fig -3 ) are a m odel o f the typical “m ethod” by Endres, who was used to prepare a prelim inary sketch, im proving it in term s o f com position and am ount o f details in subsequent versions o f the same plate W hile in other cases Reichenbach com pletely relied on E ndres’ drawings to illustrate C osta R ican orchids to be published in Xenia, the plate o f P luctuosa (Fig 1) ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at [>iip u l i N et al.: A note on Pleurothallis luctuosa (Orchidaceae) 243 Fig 3: Drawing of Pleurothallis luctuosa by A R Endres [W-Rchb Orch 36294] was ostensibly prepared from R eichenbach’s ow n sketches, based on the living plant sent by Endres and flow ered in Hamburg In 1923, in his contributions to the orchidology o f Central A m erica, Schlechter - who had exam ined E ndres’ records in V ienna - m entioned two additional collections o f P 244 ©Naturhistorisches Museum Wien, download des unter Naturhistorischen www.biologiezentrum.at Annalen Museums in Wien, B // -> Fig 4: Pleurothallis luctuosa RcHB.f (from Pupulin 5311) A: Habit; B: Flower; C: Dissected perianth; D: Column and lip, lateral view; E: Lip, spread and in natural position; F: margin of petal Drawn by D Bogarin, M Femändez and F Pupulin ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at [>ij p ij l i N et al.: A note on Pleurothallis luctuosa (Orchidaceae) 245 lu ctu osa by the brothers A lfred and K urt Brade from La Palm a (S chlechter 1923), one of the main passes through the Central Volcanic ränge connecting the central valley to ihe Caribbean in C osta R ica U nfortunately, the collections exam ined by S chlechter were eventually lost in B erlin in 1943 B y 1937, w hen Prof Arnes prepared an updated checklist o f C osta Rican O rchidaceae for S tandley ’s “Flora o f C osta R ica” , P luctuosa was still known only from a lim ited num ber o f specim ens A mes (1937) quoted a Costa Rican collection by C.H L ankester from the region o f A costa (AM ES 1178), and two specimens found by M A Brenes around San Ram on, along the C aribbean slopes o f the Tilarän m ountain ränge We w ere unable to locate the last tw o collections in A m es ’ herbarium, but G ustavo A R om ero kindly pro v id ed us w ith a m an u scrip t note by Brenes about a collection m ade by him in 1925 at “Collines en Piedades, pres S Ram on, 1025 m ” (No 38/1249 H erb B renes, A M ES) W hile L a n k e ste r‘s specim en w as recorded having “flowers m aroon, lip greenish”, the flow ers o f B renes 38/1249 w ere noted as greenish yellow, m arked w ith pale red along the veins It was not until 1976, when Luer produced the first, m odern botanical illustration o f P luctuosa for his series on m iscellaneous P leu ro th allid in ae species (L uer 1976), based on an E cuadorian specimen collected in the province o f Pichincha (L uer et al 528, SEL) In his treatm ent of the species, L uer quoted collections referable to P luctuosa also from N icaragua and Colombia (L uer 1976) L u er’s interpretation was follow ed by floristic treatm ents in the M esoamerican region (i.e., H amer 1984, G arcia C astro et al 1993, P upulin 2002, Luer 2003, C orrea et al 2004, O ssenbach et al 2005) and from South A m erica (i.e., Dodson & D odson 1980), but it is now quite clear that the specim ens quoted in these works are referable to both the true P luctuosa and to its sister, an undescribed species It is fortunate that at LBG we have plants o f both the “m orphs” to critically com pare them T a x o n o m ie t r e a t m e n t Pleurothallis luctuosa R c H B f., L in n e a : ( 7 ) - F ig = H umboldtia luctuosa (RcHB.f.) K u n t z e , Revis Gen Pl 2: 667 (1891) = Acronia luctuosa (RcHB.f.) L u e r , Monogr Syst Bot M issouri Bot Gard 103: 58 (2005) Type: C osta Rica, Alajuela, San Ram on, 1867, A.R Endres s.n [holotype: W-Rchb Orch 53791!] Herb epiphytic, caespitose, erect, small, up to 18 cm tall Roots slender, flexuous, to mm in diam eter R am icauls slender, erect, terete, 2 -9 cm long, enclosed by 1-3 tubulär, adpressed, papery, brow n bracts, -3 cm long L eaves erect, fleshy, coriaceous, subsem iterete, linear-lingulate, acute, coarsely 3-dentate, 4.2-12.8 x 0.4-0.7 cm, the cuneate-rounded base narrow ing into a petiole up to 0.15 cm long Inflorescence a lax, distichous, -6 sim ultaneously flowered racem e, 3.8-4.1 cm long including the 2.4 cm long peduncle produced at the base o f the leaf from a sm all, conduplicate, linear, acute spathe Floral bracts ovate, acute, conduplicate, m em branaceous, x mm Pedicels m m long, persistent Ovarv cylindric, m m long Flow ers w ith the sepals and the petals purple, the lip green suffused w ith purple, the colum n green, the anther greenish white D orsal sepal erect, ovate, acum inate, concave, x mm, 5-veined 246 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B / / -> Fig Distribution map of Pleurothallis luctuosa (triangles) and P neglecta (circles) in Costa Rica L ateral sepals connate into an ovate, acum inate, entire or sub-bidentulate, concave synsepal, 14 x m, 3-veined Petals ovate, acute, falcate, resting on the m argins o f the synsepal, the m argins obsoletely serrato-denticulate, x mm Lip triangular-hastate, sigmoid, acute or sub-apiculate, 3.5 x 1.5 mm, hinged to the colum n foot, the basal lobes erect; disc -carinate, the keels subobsolete, incurved, flanking the reniform glenion C olum n short, cy lin d ric-su b clav ate, m m long, the clinandrium entire, the stigma apical, reniform , bilobed A nther cap incum bent, ovate, em arginate P ollinia 2, pyriform , on an ovate, retuse viscidium Fig 6: Pleurothallis neglecta Pupulin, Bogarin & M el Fernändez (from Pupulin 7019) A: Habit; B: Flower; C: Dissected perianth; D: Column and lip, lateral view; E: Lip, spread and in natural Position; F: margin of petal Drawn by D Bogarin, M Fernändez and F Pupulin from the holotype ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at I>iii>ui IN et al.: A note on Pleurothallis luctuosa (Orchidaceae) 47 248 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B / / ’ D erivation o f name: From the Latin luctuosus, sorrow ful, in reference to the death of A R Endres, who collected the type specimen Specimens examined: C o sta R ic a [Alaiuela:! San Ramon, A R E n d r es s.n [W-Rchb Orch 53791]; A R E n d res 50 [W-Rchb Orch 44222; W-Rchb Orch 2805; W-Rchb Orch 142268: AMES 118528; W-Rchb Orch 36294, drawing; W-Rchb Orch 36294, drawing; W-Rchb Orch 33417, drawing and manuscript description]; - San Ramon, Berlin, near the turning of the unpaved road to San Mateo, Cerro Pelon, Pacific watershed of the Cordillera de Tilarän lower montane rain forest, 10°00’52”N 84°23’55”W, 1200 m, scrub Vegetation with scattered trees on exposed, windy crests, F P u p u lin 1 / & E Salas-Pupulin [JBL]; - San Ramön Angeles, road from San Ramon to La Fortuna, Angeles Norte, 10°08’29.3”N 84°28’51.2”\V, 1200 m, on crest of Continental divide, premontane wet forest, remnant woods on the right of the road, F P u p u lin 2 B & E Salas-Pupulin [JBL]; - [San Jose:] La Palma, 1400 m, A Brade & K B r a d e 1 / 121 ; Sabanillas de Acosta, 06.1928, C H L a n k e s te r 1 [AMES]; [Guanacaste:] Parque Nacional Guanacaste, Estaciön Biologica Volcän Cacao, Estaciön Pitilla, 1100 m, 24.10.1990, C C h ä v e z [CR, MO]; C o sta R ic a , without specific locality, A r ! E n d res s.n [W-Rchb Orch 2308]; C o sta R ic a , without specific locality, J B L 11663 [JBL], Distribution: To assess the real distribution ränge o f P luctuosa is m ade quite difficult by the persistent confusion w ith its sister species H amer (1984) recorded P luctuosa from N icaragua and illustrated it on the basis o f H eller 8013 [SEL], w hich corresponds to the species treated hereafter as P neglecta Even though H amer (1984) reported the species as variable, it is still know n in N icaragua from a single collection only (H amer 2001), and we therefore exclude this country from the distribution o f P luctuosa We have not seen actual m aterial o f this species from the region o f C hiriqui in Panama (D ’A rcy 1987, C orrea et al 2004), but on the basis o f its m orphologic features, the Ecuadorian specim en illustrated by L uer (1976) (L uer et al 528 [SEL]) m ay perhaps correspond to a different and still undescribed species The clone “ Sweet B ay”, which received a Certificate o f Cultural M erit by the Am erican O rchid Society in November 2008, under the nam e o f P luctuosa (K ersey 2008) is surely m isidentified; however, we have no Inform ation about the geographic origin o f this highly different taxon On the basis o f the available data, P luctuosa m ust likely be considered endem ic to Costa Rica (Fig 5) and perhaps to w estem Panama D istribution and habitat: Only know n from the Tilarän ränge in northw estem Costa Rica and, possibly, also occuring in w estern Panam a Plants have been recorded growing epiphytic in prem ontane w et forest in secondary and prim ary Vegetation Phenology: plants flow er from M ay to September Pleurothallis neglecta P u p u l i n , B o g a r i n & M e l F e r n a n d e z , s p n (F ig ) Type: Costa Rica, Alajuela: San Ram on, La Paz, Continental divide, 10°08’09.9”N 84°30’48.5”W, 1130 m, prem ontane w et forest, 19 February 2008, F Pupulin 7019, R.L D ressier & A K arrem ans [holotype: USJ; isotypes: JBL, CR, W] D iagnosis: A P leurothallide luctuosa R ch b f foliis latioribus distincte conduplicatis, sepalo postico latiore 3-nervato, petalis falcatis m arginibus erosis et labello triangulare m ulto m inore duabus carinis altis dense papillosis in disco om ato recedit Herb epiphytic, caespitose, erect, small, up to 16 cm tall Roots slender, flexuous, to m m in diam eter R am icauls slender, erect, terete, -8 cm long, enclosed by 1-3 tubulär, adpressed, papery, brow n bracts, 0.2-2.7 cm long L eaves erect, fleshy, P ijp u liN ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at et al.: A note on Pleurothallis luctuosa (Orchidaceae) Fig 7: Comparison of the flowers of (A) Pleurothallis luctuosa and of (B) P neglecta Vouchers: A - Pupulin 5305; B - Pupulin 7019 Scale bar = cm 249 250 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B //-> coriaceous, narrow ly elliptic-ovate, conduplicate, acute, retuse, 5.3-8.7 x -1 cm the rounded base narrow ing into a p etiole up to 0.15 cm long Inflorescence a lax' distichous, sim ultaneously 3-flow ered racem e, -5 cm long including the 1.4 Cni long peduncle produced at the base o f the leaf from a small, conduplicate, linear, acute spathe Floral bracts ovate, acute, conduplicate, papyraceous, x mm Pedicels long, persistent Ovary cylindric, m m long Flow ers with the sepals greenish yellow w ith purple veins, the petals brow n-purple, the lip light green Dorsal sepal erect, ovate acum inate, concave, 19 x m m , 3-veined L ateral sepals connate into an ovate, acum inate, concave, apically notched synsepal, x m m , 4-veined Petals ovate, falcate, acute, erose, x mm Lip triangular, sigm oid, the lateral lobes erect-incurved, 2.5 x 1.5 mm; disc w ith high, rounded, incurved, densely papillous keels flanking a w ell developed, ob reniform glenion C olum n short, cylindric, 1.2 m m long, the clinandrium entire, the Stigma subapical, bilobed A nther cap incum bent, ovate Pollinia , pyriform , on a rounded viscidium D erivation o f nam e: From the Latin neglectus, neglected, disregarded, alluding to the fact that botanists did not pay proper attention to this species Paratypes: C o sta R ic a Alajuela [San Carlos] about km east of Ciudad Quesada, Quebrada Marin, 10°22’ N 84°24’ W, 500 m, in shade forested stream edge and cleared slopes below the Methodist Rural Center, epiphyte on horizontal branch 10 cm diameter, 17-18.05.1968, W.C B u rg er & R G Stolze [CR]; - Alajuela: San Ramon, Angeles, Reserva Biologica Alberto Manuel Brenes, 10°13’06” N 84°36’H ” W, 800-900 m, km despues de la Estaciön, por el camino de acceso principal, bosque muy hümedo tropical transiciön a premontano, epifitas en bosque secundario a orillas del camino, 26.05.2006, D B o g a r in & Botänica Forestal Avanzada-UCR [JBL]; - Alajuela: San Ramon, Angeles, Reserva Biologica Alberto Manuel Brenes, 10°13’ N 84°35’ W, 950 m, sendero Päjaro Sombrilla, hacia el noroeste de la estaciön, M B la n co [USJ]; - Alajuela: San Ramon, camino a la Reserva Biologica Alberto Manuel Brenes, 600 m, 23.01.2001, M B la n c o [USJ]; - Alajuela: San Ramon, La Paz, C ontinental divide, 10°08’09.9” N 84°30’48.5” W, 1130 m, premontane wet forest, 19.02.2008, F Pupulin , R L Dressier and A Karremans [JBL]; - Alajuela: San Ramon, Piedades, 1025 m 05.06.1925, A M B r e n e s /1 [AMES]; - without locality data, J B L 5 [JBL]; without locality data, M B la n c o [USJ], N ic a r a g u a Rivas, 11°27’00”N 85o31’00”W, 1300 m, H e lle r 013 [SEL, drawing seen] D istrib u tio n and habitat: know n from C osta R ica and N icaragua P lants have been recorded epiphytic in secondary and prim ary forest in p rem ontane w et forest and tropical w et forest, prem ontane beit transition in Tilarän ränge, northw estem Costa Rica (Fig 5) In N icaragua it is know n from the M aderas volcano at O m etepe Island, Rivas Discussion Pleurothallis neglecta can be distinguished from P luctuosa by the distinct conduplicate leaves (vs subsem iterete), the flow ers w ith the sepals greenish yellow with purple veins, the petals brow n-purple, the lip light green to yellow ish (vs sepals and the petals purple, the lip green suffused w ith purple), triangular, sm aller (2.5 vs 3.5 m m long), the petals falcate, erose (vs obsoletely serrato-denticulate) and the callus form ed by two conspicuous, densely papillose keels (vs low, glabrous) (Fig 7) Also, it resem bles the C olom bian P leurothallis am phygia L uer & E scobar but differs in the distichous, sim u ltan eously 3-flow ered racem e (vs single flow ered), the acum inate sepals (vs narrowly, long acum inate) and the color o f the flowers, w hich are purple in P amphygia A cronia w as describ ed by Carl P resl in 1827, based on the Peruvian Acronia phalangifera In 1861, R eichenbach reduced Acronia to the synonym y o f Pleurothallis P u p u l in et a l.: ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at A note on Pleurothallis luctuosa (Orchidaceae) 251 W i t h i n Pleurothallis subgen Pleurothallis sect Pleurothallis [typyfied by Epidendrum riiscifoliiim Jacq = Pleurothallis ruscifolia (Jacq.) R Br.] L u er (1986) created subsection /[croniae w ith A p h a la n g ifera as the type, including here P leurothallis luctuosa In 1998, he divided subsection Acroniae into tw o series, Series Acroniae for those species w i t h racem ose inflorescences (including P luctuosa) and series A m phygiae for those w ith sin g le-flo w ered in flo rescen ces (L u e r 1998) The sectio n s and su b section s o f Pleurothallis subgen Pleurothallis w ere not supported as m onophyletic groups by the molecular phylogenetic analysis o f the Pleurothallidinae published by P rid g eo n et al (2001) In 2001, sp ecies o f Pleurothallis subgen Pleurothallis, including m em bers o f s u b s e c t i o n s A croniae and M acrophyllae-Fasciculatae, were segregated by S z la c h e t k o and co-w orkers ( S z la c h e t k o & M a r g o n s k a 0 , S z la c h e t k o & K u la k 2006a, 2006b) in the new genus Z osterophyllanthos S z la c h & M a r g M ore recently, L u e r (2005) proposed the resurrection o f A cronia, m erging P su b section A croniae (w ith series A croniae and A m phygiae) and P su bsection M acrophyllae-F asciculatae, n ow elevated to the sectional rank, and reducing Z osterophyllanthos to the syn on ym y o f Acronia H ow ever, recent data based on nuclear ITS and preliminary plastid sequences Information (W ils o n et al in press), support the results o f P rid g eo n et al (2001) and provide no support for the creation o f segregate genera from w ithin Pleurothallis sensu stricto (i.e the creation o f the genera Acronia and Zosterophyllanthos am ong others) According to the analysis o f W ils o n et al (in press), the genus Acronia in particular, created to include subsection M acrophyllae-Fasciculatae and the subgenus Acroniae, is not a viable concept For this reason w e favor here the description o f the new species as a member o f the genus Pleurothallis Phenology: plants flow er from M ay to July A few specim ens have been recorded in flower from July to September A cknow ledgm ents We thank Gustavo A Romero, K eeper o f the Oakes Ames Orchid H erbarium , H arvard U niversity, for kindly providing inform ation and com m ents on the Costa Rican collections preserved there Acknowledgments are extended to Ernst Vitek and Heimo Rainer at W for scanning and making available for this study the specimens and drawings by A.R Endres kept in R eichenbach’s herbarium This paper is part of the project “The life and work o f A.R Endres, orchidologist in Costa R ica”, sponsored by the VicePresidency o f Research, University o f Costa Rica The scientific services o f the Costa Rican Ministery o f Environment, Energy and Telecommunications (MINAET) and its National System o f Conservation Areas (SINAC) kindly issued the collecting perm its under w hich w ild specim ens treated in this paper were collected References A mes O., 1937: Orchidaceae - In: Hist Bot Ser 28:197-306 A no nym o us, Standley P.C (ed.): Flora of Costa Rica - Field Mus Nat 1875: Obituary: M Endres - Gard Chron., n.s., 3: 605 A., G a l d a m e s C & d e S t a p f M.S., 2004: Catälogo de las Plantas Vasculares de Panama - Panama: Smithsonian Tropical Research Institute, 599 pp C orrea DA W.G., 1987: Flora of Panama Checklist and Index - Monogr Syst Bot Missouri Bot Gard 17(1): i-xxx, -3 ; 18(2): i-ix, - r cy D o d so n C H & D o d s o n P M , 980: Orchids of Ecuador - Icon P l Trop 3: - 0 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen 252 G a r c i a C a s t r o J B , M o r a d e R e t a n a D.E & R i v a s R o s s i M E., c o m u n e s a C o s t a R ic a y P a n a m a B r e n e s ia 39-40: 93-107 Museums in Wien, B //-> 1993: L is t a d e orquideas H am er F., 1984: Orchids of Nicaragua, part - Icon Gardens, plate 1115 - Sarasota, Florida H am er F., 2001: Orchidaceae - In: S t e v e n s W.D., U l l o a U l l o a C , P o o l A & M o n t ie l O.M (eds.): Flora de Nicaragua - Monogr Syst Bot Missouri Bot Gard 85 (2): 1809-1814 P I Trop fase 12 Marie Selby Botanical M (ed.), 2008: Atlanta Judging Center Awards November 2008 - Atlanta Orch Soc Bull 49 (12): 10 K e rsey L uer L uer C.A., 1976: Miscellaneous species of Pleurothallis Selbyana (1-2): 38-201 C.A., 1986: Icones Pleurothallidinarum III Systematics of Pleurothallidinae (Orchidaceae) - Monogr Syst Bot Missouri Bot Gard 20: 1-109 L uer C.A., 1998: Icones Pleurothallidinarum XVII Systematics of subgen Pleurothallis sect Abortivae, sect Truncatae, sect Pleurothallis, subsect Acroniae, subsect Pleurothallis, subgen Dracontia, subgen Unciferia Addenda to Dracula, Lepanthes, Masdevallia, Porroglossum and Scaphosepalum (Orchidaceae) - Monogr Syst Bot Missouri Bot Gard 72: 1-121 L uer C.A., 2003: Pleurothallis - In: H a m m e l B.E., G r a y u m M.H., H e r r e r a C & Z a m o r a N (eds.): Manual de plantas de Costa Rica, vol III - Missouri Botanical Garden Press: 386-452 L uer C.A., 2005: Icones Pleurothallidinarum XXVII Diyadella and Acronia sect Macrophyllaceae-Fasciculatae - Monogr Syst Bot Missouri Bot Gard 103: 1-311 O C., P u p u l i n F & D r e s s l e r R.L., 2005: Orquideas del istmo centroamericano, catälogo y estado de conservaciön Orchids of the Central American isthmus, checklist and Conservation status - Editorial 25 de mayo 243 pp - San Jose sse n b a c h C., P u p u l i n F & J e n n y R., 2010: Orchid itineraries o f Augustus R Endres in Central America: a biographic and geographic sketch - Lankesteriana 10(1): 19-47 O ssen ba c h A.M., S o l a n o R & C h a s e M.W., 2001: Phylogenetic relationships in Pleurothalli dinae (Orchidaceae) Combined evidence from nuclear and plastid DNA sequences Amer J Bot 88 (12): 2286-2308 P r id g e o n F., 2002: Catälogo revisado y actualizado de las Orchidaceae de Costa Rica Lankesteriana 4: 1-88 P u p u l in R e ic h e n b a c h H.G., 1875: Masdevallia chimaera - Gard Chron., n.s., 4: 258 R e ic h e n b a c h H.G., 1877: Pleurothallis (Brachystachyae) luctuosa - Linnaea 41: 48 R e ic h e n b a c h H.G., 1878: Pleurothallis luctuosa - Xenia Orch 3: 15-16, pl 210 R., 1923: Beiträge zur Orchideenkunde von Zentralamerika II Additamenta ad Orchideologiam Costaricensem - Repert Sp Nov Regni Veg Beih 19 S chlechter D.L & M a r g o n s k a B.H., 2001: Genera et species Orchidalium - Polish Bot J 46: 113-121 Szlachetko S z l a c h e t k o D L & K u l a k M , 2006a: New combinations in the genus Z o ste ro p h ylla n th o s S z l a c h & M a r g (Orchidaceae, Pleurothallidinae) - Richardiana 6: 131-135 S z l a c h e t k o D.L & K u l a k M , 2006b: New combinations in the genus Z o ste ro p h ylla n th o s S z l a c h & M a r g (Orchidaceae, Pleurothallidinae) - Richardiana 6: 183-195 W M., B e l l e C., D a n g A., H a n n a n P., K e n y o n C., Low H , S t a y t o n T & W o o l l e y M (in press): A phylogenetic analysis of the genus Pleurothallis, with emphasis on Pleurothallis subsection Macrophyllae-Fasciculatae, using nuclear ITS and chloroplast DNA sequencing — Proc 3rd Scientific Conference on Andean Orchids Quito Pontificia Universidad Catölica del Ecuador, 2008 il s o n ...240 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B ] p on the crucial facts o f his biography... from Xenia Orchidacea, 1878 242 ©Naturhistorisches Museum Wien, download unter Annalen des www.biologiezentrum.at Naturhistorischen M useums in Wien, B / /-> v• Fig 2: Drawing of Pleurothallis... additional collections o f P 244 ©Naturhistorisches Museum Wien, download des unter Naturhistorischen www.biologiezentrum.at Annalen Museums in Wien, B // -> Fig 4: Pleurothallis luctuosa RcHB.f