Selbyana 25(2): 182-209. 2005.

A MONOGRAPH OF ALLOPLECTUS ()

JOHN L. CLARK Department of Botany, MRC-166, U.S. National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013-7012; Current address: Department of Biological Sciences, George Washington University, 2023 G Street, NW, Washington, DC 20052 USA. Email: [email protected]

ABSTRACT. Alloplectus (Gesneriaceae-Episcieae), whose range extends from Costa Rica to the Andes of Colombia, Ecuador, and Peru, comprises five species. The are obligate epiphytes with tubular non­ resupinate flowers. The generic circumscription of Alloplectus is based on phylogenetic analyses of mor­ phological and molecular data from the ribosomal (nrDNA) internal transcribed spacer region (ITS) and the intergenic (tmH-psbA) chloroplast spacer region (details of the analyses are published separately). Alloplectus has a confusing taxonomic history with more than 150 names attributed to this poorly defined group that until recently had not been phylogenetic ally evaluated with material representing the generic type species, Alloplectus hispidus (Kunth) Mart. Many of the species traditionally recognized as Alloplectus are here transferred to , Nematanthus, Drymonia, Glossoloma, and Crantzia, of which the latter two are resurrected generic names to accommodate well-supported clades. Replacing the traditional poly­ phyletic concept of Alloplectus with a monophyletic Alloplectus presented here requires 29 new combi­ nations, two lectotypifications, and six neotypifications. A nomenclator of all excluded or uncertain species names attributed to Alloplectus is included with currently accepted names.

Key words: Gesneriaceae, , Alloplectus, Glossoloma, Crantzia, Columnea, Drymonia

INTRODUCTION come a collection of species that lacked unifying synapomorphies. The genus Alloplectus Mart. has been a catch­ The taxonomic treatment here recognizes the all group for taxa that do not fit into other genera genus Alloplectus as a well-supported clade (bs in the tribe Episcieae. The traditionally recog­ = 100% in Clark et al. in press) united by the nized species of Alloplectus constitute a poly­ following unambiguous morphological synapo­ phyletic group of 40 species that requires the morphies: indistinguishable dry secondary leaf transfer of 29 names from Alloplectus into Ne­ venation, glabrous internal corolla surface, and matanthus (1 sp.) and Drymonia (4 spp.) and corollas constricted apically. Other characters of recognition of Glossoloma (21 spp.) and Crant­ Alloplectus that are useful in distinguishing it zia (3 spp.) for two well-supported clades. Thus, from Glossoloma are the subwoody perennial epiphytic habit, non-resupinate flowers, and pro­ the revised Alloplectus is here treated as a genus fusely branched stems. Characters that are help­ of five species. This paper provides a taxonomic ful for differentiating Alloplectus from other revision of Alloplectus based on recent phylo­ closely related genera of Episcieae are listed in genetic analyses of molecular and morphologi­ TABLE 1. cal data (Clark et al. in press). The five accepted species of Alloplectus range Most characters traditionally used to define from Costa Rica to Peru and are most abundant Alloplectus, such as a fleshy bivalved dehiscent in the northern Andes in Colombia. Many of the capsule, a pendent inflorescence of a reduced former species of Alloplectus, which form a pair-flowered cyme, and a haploid chromosome well-supported clade now recognized as the ge­ number of N = 9, are symplesiomorphic char­ nus Glossoloma, are easily differentiated in the acters shared with other genera such as Dry­ field by the presence of resupinate flowers. The monia and Paradrymonia. Other characters pre­ majority of species traditionally recognized in viously used to define Alloplectus, such as a tu­ Alloplectus will be treated in a revision of Glos­ bular or pouched corolla tube, are convergent soloma (Clark unpubl. data). with Nematanthus. To a certain extent, Alloplec­ tus was "defined" by what it lacked. For ex­ PHYLOGENY ample, species with poricidal anther dehiscence were treated as Drymonia, species with translu­ The neotropical Gesneriaceae comprise mem­ cent berries were treated as Corytoplectus, spe­ bers of two subfamilies, Gesnerioideae and part cies with non-translucent berries were treated as of Coronantheroideae (Burtt & Wiehler 1995). Columnea, and species with stolons were treated The subfamily Coronantheroideae contains nine as Alsobia or Episcia. Thus Alloplectus had be- genera and 20 species and extends from Chile 182 CLARK: ALLOPLECTUS MONOGRAPH 183

TABLE 1. Comparison of Alloplectus, Glossoloma, Coiumnea, Drymonia, and Crantzia,

Characteristic Alloplectus Glossoloma Columnea Drymonia Crantzia Leaves Isophyllous Isophyllous Anisophy lIous Isophyllous Isophyllous (rarely isophyl- lous) Flower orienta- Non-resupinate Resupinate Non-resupinate Non-resupinate Resupinate or tion non-resupi- nate Habit Obligate epi- Terrestrial Facultative epi- Epiphytic or ter- Epiphytic or phyte (rarely epi- phyte or terres- restrial terrestrial phytic) trial Fruit Fleshy capsule Fleshy capsule Berry (rarely Fleshy capsule Fleshy capsule fleshy capsule) (rarely fleshy berry) Anther dehis- Longitudinal Longitudinal Longitudinal Poricidal and Longitudinal cence longitutinal Corolla pouch Terminal when Terminal when Medial when Medial when Terminal when present present present present present

to the South Pacific Islands and Australia (Wieh­ well supported (bs = 100% in Clark et al. in ler 1983), The Gesnerioideae contains five press) by both molecular and morphological tribes, 56 genera, and more than 1800 species data. The molecular data are from the ribosomal and represents nearly half the worldwide diver­ (nrDNA) internal transcribed spacer region sity of the family Gesneriaceae (Wiehler (ITS), a noncoding region between 18S and 26S 1983, Burtt & Wiehler 1995). The Episcieae is of the nrDNA and the intergenic spacer, trnH­ a member of the Gesnerioideae and ranks as the psbA of the cpDNA. Details of the ITS analysis most diverse tribe in the family with 22 genera are described by Clark and Zimmer (2003) and and an estimated 784 species, or roughly 21 % by Clark et al. (in press). of all Gesneriaceae. Episcieae is also the least The phylogenetic analysis (Clark et al. in studied, and generic concepts remain poorly de­ press) samples 21 of the 22 traditionally recog­ fined, partly because of a simplistic use of fruit nized genera with 155 species and 16 generic structure by earlier workers in delimiting genera. type species. To date, it is the most extensive A special need exists for more generic level phy­ taxon sampling within Episcieae. Although still logenetic analyses with greater sampling of spe­ greater taxon and character sampling is needed, cies diversity. the analysis is adequate to provide a framework Of the 22 genera in Episcieae, CoZumnea L. for the revision of AZZopZectus. The most recent is the only genus that has been consistently taxonomic treatment of AlZoplectus is well over shown to be monophyletic (with the inclusion 100 years old (Hanstein 1865). Hanstein's pub­ here of CoZumnea dielsii Mansf.) using morpho­ lication focused on the Gesneriaceae at the bo­ logical and molecular data (Smith 1994, Smith tanical garden in Berlin and provided an over­ & Sytsma 1994a, 1994b, 1994c). The lack of view of the family. Alloplectus was only a part monophyly for large genera (i.e., >20 spp.) in of Hanstein's (1865) entire monograph of the Episcieae has been suggested for Drymonia Gesneriaceae known at that time, but he recog­ Mart. (Smith 2000, Clark & Zimmer 2003), Epi­ nized 30 species of AZZopZectus in his treatment. scia Mart. (Smith et al. 1997), AlZoplectus Mart. More recent estimates range from 60 (Skog (Clark & Zimmer 2003), Paradrymonia Hanst. 1978 [1979]) to 75 species (Burtt & Wiehler (Smith & Carroll 1997, Smith 2000, Clark & 1995). Zimmer 2003), Codonanthe (Mart.) Hanst. Various sister taxa relationships for AlZoplec­ (Clark & Zimmer 2003), and Nematanthus tus have been proposed in the literature (Smith Schrad. (Clark & Zimmer 2003). NautiZocaZyx & Sytsma 1994a, 1994b, 1994c; Smith & Car­ Hanst. has 70-80 species (Burtt & Wiehler roll 1997; Smith 2000; Zimmer et al. 2002). 1995) and is one of the few large (i.e., >20 spp.) Only one publication, however, included the episcioid genera that has not been tested ade­ type species, AZZoplectus hispidus (Clark & Zim­ quately. mer 2003). Previous analyses were focused on The recognition of AZZopZectus is based on a resolving tribal-level phylogeny of the family total evidence analysis of two molecular datasets (FIGURE 1). The plastid markers such as ndhF, for 155 species and .a morphological matrix of trnL-F, and trnE-T used in previous analyses 120 species and 99 characters. AlZoplectus is were appropriate for addressing tribal level phy- 184 SELBYANA Volume 25(2) 2005

- =80-100% Bootstrap values only one previous analysis (Clark & Zimmer = 60-80% Bootstrap values 2003) included a species that represents the cur­ = <60% Bootstrap values rent clade-based concept of Allopleetus. Previ­ ous studies (FIGURE lA-E) can be summarized "..... Glossoloma I"'" Glossoloma r<]ColUmnta as follows. FIGURE lA. Drymonia and Glossoloma (as :, .. ~. <] ! .. ~ /I "Allopleetus") form a clade that is sister to Col­ .• :. , Drymonia Drymonia ~ :"""'-J Columnta .. ,L<] umnea. Sequence data from the chloroplast gene ndhF (Smith & Carroll 1997, Smith et al. 1997) L .... Columnea ...... GlossoWma and a combined analyses of the nr-ITS region

names were never formally transferred to Allo­ (Schott) Wiehler. Oersted created the new genus plectus. Prionoplectus, including, for example, Crantzia Three names were subsequently transferred cristata (L.) Scop., for species with ovate sepals from other genera (Besleria cristata L., B. his­ with cordate bases. Initially, Hanstein (1854) pida Kunth, and B. coccinea AubI.) and given also recognized Alloplectus Mart., but restricted new combinations (Alloplectus cristatus, A. his­ his generic circumscription to species with pidus, and A. coccineus) in the Index (Martius pouched corollas (e.g., Alloplectus circinatus 1832). Thus, the 1832 Index validated three Mart., which is now Drymonia coccinea (AubI.) names making Alloplectus a genus of five spe­ Wiehler; Alloplectus speciosus Poepp., which is cies. now Corytoplectus speciosus (Poepp.) Wiehler; The 1905 International Code of Botanical No­ and Alloplectus concolor Hook., which is now menclature (ICBN) conserved the name Allo­ Nematanthus lanceolatus (Poir.) Chautems. plectus Mart. (Martius 1829) over Crantzia Hanstein published a new genus, Glossoloma, Scop. (Scopoli 1777) and Vireya Raf. (Rafin­ for species with tubular and ventricose corollas esque 1814). Green (1929) selected Alloplectus (e.g., Glossoloma tetragonum Hanst.). Approx­ sparsiflorus Mart. as the type species for Allo­ imately a decade later, Hanstein (1865) signifi­ plectus because it was a widely distributed spe­ cantly modified his initial classification from cies. Green's typification was accepted and cited 1854 and decided to combine many of the Oer­ in subsequent Codes. Notes pertaining to the se­ sted genera and five of his own genera into Al­ lection of A. sparsiflorus and typification are not loplectus. Hanstein further divided Alloplectus included in the published proposal (Green into the following five subgenera: Calanthus, 1929), but an explanation of Green's notes can Erythranthus, Heintzia, Glossoloma, and Eual­ be found in a mimeographed foolscap-size cir­ loplectus. He also recognized three sections of cular (Stafleu 1956). Wiehler (1971 [1972]) not­ the typical subgenus Eualloplectus and included ed that A. sparsiflorus was a synonym of Ne­ a section of dubious species that he could not matanthus hirtellus (Schott) Wiehler, leaving Al­ assign to any of the subgenera in his classifica­ loplectus without a type and only two remaining tion. In total, Hanstein treated 29 species of Al­ species from Martius's generic description. The loplectus and until the present study, his was the proposal by Feuillet and Skog (1990) to con­ most comprehensive monographic revision un­ serve A. hispidus (Kunth) Mart. as the type spe­ dertaken for Alloplectus. cies for the genus was accepted. Their proposal Kuntze (1891) transferred all of the species of considered the remaining two species originally Alloplectus into Columnea. Because these two attributed to Alloplectus by Martius. Feuillet and genera had traditionally been kept apart, there Skog (1990) chose A. hispidus over A. cristatus, was an extensive section in the Kuntze paper because the fruit of the latter had been reported listing new names to avoid homonyms as a re­ to be a berry. Alloplectus hispidus was known sult of all of the new combinations. Kuntze also to have fleshy capsules, thus making it more rep­ recognized Crantzia as the older and valid name resentative of the generic description given by for Alloplectus, but it was not pertinent because Martius. It should be noted, however, that recent he combined all the species into Columnea. field observations by the author on the islands Fritsch (1893-1894) maintained Columnea of Martinique and Dominica have shown that separate from Alloplectus, but he used the older the fruit of A. cristatus is not a berry, but instead name Crantzia. Fritsch recognized six sections a bivalved capsule, typical of Alloplectus. as part of Crantzia: Calanthus Oerst. ex Hanst., Two botanists, Oersted (1816---1872) and Han­ Macrochlamys Decne., Heintzia Karst., Glosso­ stein (1822-1880), who were developing a clas­ loma Hanst., Alloplectus Mart., and Trichantho­ sification of Gesneriaceae during the same time des Kuntze. The Fritsch classification was sim­ period, based their classifications on extensive ilar to that of Hanstein and he frequently re­ studies of herbarium and living material. Oersted ferred to Hanstein's subgeneric classification. traveled throughout Central America (1845- More recently, Alloplectus and Columnea 1848), and Hanstein had access to an extensive have been treated as separate genera by recog­ living collection at the botanical garden in Ber­ nizing the presence of berries in the latter as an lin. Oersted (1858, 1861) recognized Alloplectus important defining generic character (Morton Mart., but restricted it to species with plane 1963, Wiehler 1973, Skog 1978 [1979], Wiehler (non-cucullate) sepals, such as Alloplectus sem­ 1983, Kvist & Skog 1993, Smith 1994). The icordatus Poepp., which is now Drymonia sem­ documentation of poricidal anthers shedding icordata (Poepp.) Wiehler; Alloplectus cocci­ powdery pollen grains by Moore (1955) and fur­ neus (AubI.) Mart., which is now Drymonia coc­ ther elaborated by Wiehler (1983) as a generic cinea (AubI.) Wiehler; and Alloplectus sparsiflo­ character in Drymonia has helped differentiate rus Mart., which is now Nematanthus hirtellus many of the fleshy capsuled species of Drymon- 186 SELBYANA Volume 25(2) 2005

ia from Alloplectus. Although poricidal anthers TABLE 2. Index to Alloplectus collections examined, are a synapomorphy for Drymonia, they have listed alphabetically by collector name(s). Collec- been lost in several clades within the genus tion numbers are included along with species de- (Clark et al. in press). The lack of poricidal an­ termination. thers in Drymonia is probably correlated with a Collector(s) Collection # Species switch in pollination syndromes from Euglossine bees to birds. Alvarez, A. & M. Tirado 1529 A. hispidus Andre, E. 1514 A. hispidus 2045 A. hispidus TAXONOMIC TREATMENT Araya, E, et al. 264 A. weirii Alloplectus Mart., Nov. Gen. Sp. PI. 3: 53. 1829. Ariste-Joseph, Bro. S.n. A. hispidus CONSERVED TYPE: Besleria hispida Kunth Asplund, E. 8072 A. hispidus Barclay, A., et al. 3395 A. hispidus [=Alloplectus hispidus (Kunth) Mart.] (see Benavides, 0., et a1. 4365 A. hispidus Feuillet & Skog 1990). Besse, L., et a1. 1155 A. weirii Subshrubs always epiphytic; stems terete to 1161 A. weirii quadrangular, horizontal, usually scandent. Betancur, J., et al. 7022 A. aquatiUs Bohlin, J.E., et al. 1032 A. hispidus Leaves opposite, nearly equal in a pair, petiolate, Bonpland, A. 1776 A. hispidus elliptic to ovate or oblanceolate, entire to serrate, 2104 A. hispidus secondary venation inconspicuous when dry. Bj1Igh, A. 86568 A. hispidus Flowers axillary, solitary to fascicles of 2 to 3, Camargo, G., L.A. 7437 A. hispidus bracteate, peduncles rarely evident; calyx usu­ Campos, J. & S. Corrales 3441 A. weirii ally colored or green, the 5 lobes nearly free, Carvajal, A. 166 A. weirii broadly ovate, subequal, entire to strongly ser­ 368 A. weirii rate, pubescent to pilose; corolla tubular and Cer6n M., C.E. & 38680 A. weirii usually ampliate, rarely hypocyrtoid, yellow or T. Davila Cer6n M., C.E. & 32721 A. weirii red, oblique to horizontal in the calyx, constrict­ C. Montalvo A. ed in the throat, gibbous at the base, often ven­ Clark, J.L. 2307 A. weirii tricose below the mouth, the 5 lobes of the limb Clark, J.L. & H.E 5788 A. weirii usually small; stamens 4, not exserted, the fila­ Greeney 5792 A. weirii ments adnate to the base of the corolla tube, and Clark, J.L. & J. Rea 8016 A. weirii forming an open sheath, the anthers transversely Clark, J.L. & S. Yustiz 6871 A. aquatilis oblong, coherent or becoming free, dehiscing by 6875 A. aquatiUs longitudinal slits; nectary disc a bilobed gland, Clark, J.L., et al. 5625 A. hispidus 5701 A. weirii rarely single-lobed; ovary superior, pilose, the 7720 A. hispidus style included. Fruit a fleshy bivalved capsule 7726 A. hispidus surrounded by the persistent calyx; seeds nu­ 7744 A. hispidus merous, longitudinally striate. 8210 A. tessmannii 8914 A. weirii Croat, T.B. KEy TO THE SPECIES OF ALLOPLECTUS 78842 A. weirii Cuatrecasas, J. 8752 A. weirii 1. Calyx lobe margin fimbriate .. Alloplectus weirii 19304 A. hispidus 1'. Calyx lobe margins serrate to entire. 23333 A. inflatus 2. Corolla red 23465 A. hispidus 3. Corolla tubular with medial pouch. Pedi- 23484 A. injlatus cels 0.5-1 cm long ...... Daniel, Bro. 3444 A. hispidus ...... Alloplectus tessmannii 3445 A. hispidus 3'. Corolla tubular with terminal pouch (i.e., 4471 A. weirii hypocyrtoid). Pedicels 2-7 cm long .... Diaz, C. & H. Beltran 3341 A. weirii ...... Alloplectus injlatus Diaz, e, S. 2677 A. injlatus 2'. Corolla yellow Diaz, e, S., et al. 2826 A. hispidus 4. Calyx lobe margins valvate to spread­ Duque-Jaramillo, J.M. 3454 A. hispidus ing; stems sparingly branched .... Dwyer, J.D. & 9581 A. weirii ...... Alloplectus aquatilis B. Macbryde 4'. Calyx lobe margins connivent to over­ Escobar, L.A. & 6216 A. hispidus lapping; stems prolifically branched .. B.A. Stein ...... Alloplectus hispidus Escobar, L.A. & A. Uribe 3621 A. weirii Fernandez A., J.L., et a1. 14870 A. hispidus 14881 A. hispidus ALLOPLECTUS COLLECTIONS EXAMINED Fonnegra, R., et al. 3479 A. hispidus The Alloplectus collections examined are list­ Fosberg, ER. 20121 A. hispidus 20862 A. inflatus ed in TABLE 2. The table comprises an index of CLARK: ALLOPLECTUS MONOGRAPH 187

TABLE 2. Continued. TABLE 2. Continued.

Collcctor(s) Collection # Species Collcctor(s) Collection # Species Garcia, D. 189 A. weirh 011gaard, B., et al. 35762 A. weirii Garcia Barriga, H. 17203 A. hispidus 58010 A. hispidus Garcia Barriga, H. & 20261 A. infiatus Obando, N., et al. 167 A. weirii R. Jaramillo Mejia 20545 A. weirii Palacios, W. 2573 A. hispidus Gentry, A., et a1. 61191 A. weirii 5936 A. weirii Gomez, L.D., et al. 20789 A. weirh 6351 A. weirh Goudot, J. s.n. A. hispidus Palacios, W. & 3678 A. hispidus 3 A. hispidus H. van der Werff Grant, M.L. 9184 A. weirii Pearce, R. S.n. A. hispidus 9522 A. infiatus Pedersen, H.B. & 104349 A. hispidus 9596 A. i11;f/atus P Lozano Grubb, PJ., et a1. 1271 A. weirii Pederson, H.B., et a!. 104445 A. hispidus Haber, W.A. 852 A. weirii Pennell, F. W. 2496 A. hispidus 996 A. weirii 2647 A. hispidus 1007 A. weirii 9081 A. weirii 11298 A. weirii 9383 A. injlatus Harling, G. & 21900 A. hispidus Pennell, F.w. & 5782 A. weirii L. Andersson E.P Killip Harling. G., et al. 8204 A. hispidus Penuela, L. & 16 A. infiatus Hawkes, J.G. & 23 A. infiatus J. C1avijo H. Garcia Barriga Poortmann, H. 143 A. hispidus Herrera, G. 4924 A. weirii Prieto, F. 212 A. hispidus Holm-Nielsen, L., et a1. 6969 A. weirii Ramirez P., B.R. 5365 A. hispidus Huertas, G. & 5007 A. infiatus Ramirez P, B.R. & 3954 A. hispidus L.A. Comargo J.A. Cuayal M. 4892 A. hispidus Humboldt, A. & 1878 A. hispidus Ramos, J.E., et al. 2574 A. weirii A. Bonpland Romero-Castaneda, R. 937 A. hispidus Idrobo, I.M., et al. 3376 A. hispidus Ruiz Teran, L. & 8912 A. hispidus 3837 A. hispidus M. Lopez E A. weirh Jaramillo M., R., et al. 6026 A. hispidus Rusby, H.H. 2424 A. hispidus Jaramillo, J. 5314 A. hispidus Rusby, H.H. & 786 Jorgensen, PM., et a1. 1374 A. hispidus EW. Pennell 893 A. hi'lJidus Rusby, H.H., et a1. 15335 A. hispidus Killip, E.P 5578 A. weirii 16774 A. hispidus 5926 A. weirii Sagastegui, A., et a1. Sanchez Vega, 1. & 8477 A. tessmannii Killip, E.P & H. Garcia 33806 A. hispidus M.O. Dillon 8653 A. tessmannii Killip, E.P & G. Varela 34612 A. infiatus Schlim, L. 1089 A. weir;; Kirkbride, J.H. & 4009 A. weirii Schneider, M. 686 A. hispidus H. Chambra R. 784 A. hispidus K vist, L.P, et a1. 60393 A. weirii 1196 A. infiatus Lawrance, A.E. 357 A. weirii Schunke v., J. 9143 A. weirii Leiva G., S., et a1. 1574 A. weirii Scolnik, R., et al. 247 A. hispidus Lellinger, D.B. & 1253 A. hispidus Silva, H. 272 A. hispidus .T.J. White Silverstone Sopkin, 1527 A. hispidus 309 A. tessmannii Leo, M. EA., et a1. Liesner, R.L. & 14694 A. weirii Smith, D.N. & 4723 A. weirii E. Judziewicz S. Vasquez S. A. hispidus Londono, X. & 90 Smith, J.F. & C. Diaz 1655 A. weirii A. weirii L.P Kvist 809 Sodiro, L. s.n. A. hi,pidus A. hispidus Lozano, C., G. 2172 Spruce, R. 5179 A. hispidus 3492 A. hispidus Stevens, W.D. 13913 A. weirii 4063 A. infiatus Steyermark, J.A. 54699 A. weirii Luer, c., et al. 7566 A. weirii 54882 A. weirh Lugo S., H. 1124 A. weirii 56528 A. aquatilis 1J40 A. weirii 57247 A. hispidus Marin, F. 2116 A. weirii 91523 A. hispidus Meier, W., et a1. 4197 A. hispidus 98948 A. hispidus 8471 A. hispidus 98977 A. hispidus Murcia, e. 12 A. infiatus Steyermark, I.A. & 97648 A. hispidus Mutis, J.e. 2249 A. weirii G. Bunting 0llgaard, B. & 90304 A. hispidus Steyermark, J.A. & 111158 A. hispidus B. Eriksen V. e. Espinosa 188 SELBYANA Volume 25(2) 2005

TABLE 2. Continued. densely hirsute, pedicel glands absent, flower posture horizontal at anthesis. Calyx lobes 5, Collector(s) Collection # Species nearly free, plane, margins valvate to completely Steyermark, I.A., et al. 98427 A. hispidus separate, erect, equal, broadly ovate to lanceo­ 100658 A. aquatilis late, 20-25 mm long, 5-10 mm wide, base trun­ 114307 A. hispidus cate, apex attenuate, margin serrate, pale green, Suin, L., et al. 1805 A. weirii outside densely hirsute, inside densely hirsute. Tessmann, G. 1800 A. tessmannii Corolla 2.5-3.5 cm long, tubular, tube to 3.5 cm Tracey, 1.A. 329 A. hispidus long, gibbous basally on upper surface, spur ab­ Triana, J.J. s.n. A. hispidus sent, long axis of corolla oblique relative to ca­ 607 A. hispidus lyx; base 3-5 mm in diam., middle ampliate, 2454 A. weirii 2484 A. weirii becoming apically ventricose on lower surface, 2499 A. hispidus throat constricted, not appearing laterally com­ 4163 A. hispidus pressed, to 3 mm (at mouth), outside sparsely 4163 A. weirii hirsute becoming densely hirsute distally, inside Dribe D., L. 1016 A. hispidus glabrous, interior spotting absent, limb reflexed, 1758 A. weirii subregular, 3 mm wide, yellow, lobes nearly 2594 A. infiatus equal, rotund, ca. 1 mm long, 1.5 mm wide, 2961 A. hispidus rounded, entire. Nectary gland bilobed, on dor­ 3195 A. hispidus sal surface of ovary, glabrous. coiling 3535 A. infiatus Filaments 4915 A. hispidus after anthesis, to l.4 cm long (free portion), ad­ 5568 A. hispidus nate to base of corolla tube for 3 mm, connate 6387 A. infiatus for 1 mm, forming an open sheath, glabrous; an­ Vargas, W. G. 3224 A. weirii thers broader than long, ca. 2 mm long, ca. 3 3980 A. hispidus mm wide, dehiscing by longitudinal slits; stam­ Vivar C., F. & 3733 A. hispidus in ode absent. Ovary ovoid, ca. 6 mm long, ca. O. Proafio 5 mm wide, pilose, style 1.1 cm long, glabrous, Weaver, R.E. & 1567 A. infiatus stigma stomatomorphic. Fruit a fleshy capsule, H. Kennedy pendent when ripe, ovoid when immature, then Weir, I. 217 A. weirii Werff, H. van der & 9620 A. hispidus becoming bivalved when mature, 1.4 cm long, W. Palacios 1.0 cm wide, pilose. Seeds numerous, ellipsoid, Y:inez, A.P. & 91 A. hispidus 1.2 mm long, 0.2 mm wide, longitudinally stri­ M. Gavilanes ate, light red. PHENOLOGY: Collected in flower during Janu­ ary, February, and May; in fruit in October. collector name(s) listed alphabetically along with collection numbers and species names. DISTRIBUTION: Colombia (Cundinamarca) and Alloplectus aquatilis C.V.Morton, Gesneriaceae, Venezuela (Merida and Tachira); eastern An­ in J.A. Steyermark, Fieldiana, Bot. 28: 521. dean slopes in wet montane forest; 2400-2900 1953. TYPE: Venezuela-Me:rida, near wa­ m (FIGURE 2). terfall between El Molino and ridge above REPRESENTATIVE SPECIMENS: Colombia-Cun­ San Isidro Alto, 2430-2895 m, 14 May dinamarca: Regi6n del Sumapaz, Carretera San­ 1944, J.A. Steyermark 56528 (holotype: ta Rosa-Santo Domingo, Betancur et al. 7022 US!; isotypes: F! NY!). (COL, US). Venezuela-Merida: Dirt road be­ Lianas epiphytic, stems scandent, frequently tween Torondoy and Pico Aguila, entrance to branched; 2-3 m long, 1-5 mm in diam., woody, Parque Nacional Culata. Clark & Yustiz 6871 internodes 3-8 cm long, terete, glabrescent be­ (MER, NY, PORT, SEL, US, VEN); Clark & low, hirsute with yellowish hairs above, leaf Yustiz 6875 (MER, NY, PORT, SEL, US, scars flush with stem surface. Leaves opposite, VEN).-Tachira: Quebrada las Copas, Steyer­ equal or subequal in a pair; petioles 1-2 cm mark et al. 100658 (NY, US, VEN). long, densely hirsute; blades membranous when Alloplectus aquatiUs is distinguished from dry, ovate, 6-7 cm long, 2.5-4.0 cm wide, base other members of the genus by calyx lobes that obtuse to rounded, apex acuminate, margin are broadly ovate and spreading. It is similar to bluntly serrate to crenate, adaxially light green, A. hispidus, but differs in having calyx lobes that uniformly hirsute, abaxially pale green, hirsute are completely separate compared to the conni­ on veins, lateral veins 4-5 per side. Flowers sol­ vent to valvate margins of A. hispidus (FIGURE itary, bracteoles not observed; pedicels equal to 3B). It is also differentiated by membranous or slightly longer than petiole, 2-3 cm long, leaves that are sparingly pilose in contrast to the CLARK: ALLOPLECTUS MONOGRAPH 189

5° 50

A Alloplectus aquatilis Alloplectus hispidus 00 00 •0 Alloplectus inflatus * Alloplectus tessmannii 0 Alloplectus weirii

50 5D

0 500 lOOO KM

10° 10°

FIGURE 2. Distribution of Alloplectus aquatiUs C.Y.Morton, A. hispidus (Kunth) Mart., A. injlatus J.L.Clark & L.E.Skog, A. tessmannii Mansi., and A. weirii (Kuntze) Wiehler. This map was generated using ArcView GIS 3.2a (ESRI 2000). coriaceous leaves with densely pilose trichomes wide, base acute to obtuse, apex acute, margin of A. hispidus. serrate-crenate, adaxially dark green, densely pi­ lose-hispid, abaxially light green, densely pi­ Alloplectus hispidus (Kunth) Mart., Nov. Gen. lose-hispid, lateral veins 3-5 per side. Flowers Sp. PI. 3: 57. 1829. Besleria hispida Kunth solitary, bracteoles not observed; pedicels longer in H.B.K., Nov. Gen. Sp. PI. 2: 320 (folio), than the petiole, 1.5-3.0 cm long, densely pi­ 399 (quarto). 1817 [1824]. Columnea hum­ lose-hispid, pedicel glands absent, flower pos­ boldtiana Kuntze, Revis. Gen. PI. 2: 471. ture horizontal to erect at anthesis. Calyx lobes 1891, non Columnea hispida Swartz. TYPE: 5, nearly free, plane, erect, 4 subequal, broadly Colombia-Narifio, between city of Pasto ovate, 10-15 mm long, 4-6 mm wide, base and Mamendoy, valley of Rio Mayo, 1900 truncate, apex acute, margin laciniate to serrate, m, Nov 1801, A. Bonpland 2104 (holotype: light green, outside densely pilose, inside spar­ P!; isotypes: B-W (photo: US!), P-2 sheets!; ingly pilose; fifth (ventral) lobe smaller and nar­ fragment: F!). FIGURE 3. rower, broadly oblong, ca. 1.5 cm long, ca. 0.5 Vines epiphytic, stems branched, climbing or cm wide. Corolla 2.5-3.5 cm long, tubular, tube sprawling, adhering to other plants by adventi­ 2.2-3.3 cm long, gibbous basally on upper sur­ tious roots; growing 2-5 m above ground, 5-8 face, spur absent, long axis of corolla oblique mm in diam., woody, internodes 2-6 cm long, relative to caJyx; base 4-5 mm in diam., middle terete, glabrescent below, densely pilose above, ampliate, becoming apically ventricose on lower leaf scars flush with stem surface, nectary glands surface, throat constricted, not appearing later­ at base of petiole present. Leaves opposite, equal ally compressed, ca. 0.5 cm wide (at mouth), or subequal in a pair; petioles 1.3-1.8 cm long, outside densely pilose-hirsute, inside glabrous, densely pilose; blades coriaceous when dry, interior spotting absent, limb reflexed, zygomor­ ovate-elliptic, 3.0-6.5 cm long, 2.0-3.0 cm phic, ca. 0.6 cm wide, yellow, lobes nearly 190 SELBYANA Volume 25(2) 2005

FIGURE 3. Alloplectus hispidus (Kunth) Mart. A. Habit. B. Lateral view of flower. C. Lateral view of mature fruit with persistent calyx. D. Front view of flower. Photographic images of field collection 1.L. Clark, L. lost & M. Mailloux 7720 (US). equal, rotund, reflexed, ca. 2 mm long, ca. 3 mm REPRESENTATIVE SPECIMENS: Colombia-An­ wide, rounded, entire. Nectary gland bilobed, on tioquia: 10 km E of Sonson, Scolnik et at. 247 ventral surface of ovary, glabrous. Filaments (US); Mpio. Frontino, Correg. Nutibara, Alto de curved or twisted after anthesis, ca. 1-2 cm (free Cuevas, Quebrada del Oso, Fonnegra et at. 3479 portion) long, adnate to base of corolla tube for (US); Mpio. de Caldas, Vereda La Corrala, road ca. 2 mm, connate for I mm, forming an open between Finca "La Zarza" and Alto del Gallin­ sheath, glabrous; anthers broader than long, ca. azo, Escobar & Stein 6216 (COL, NY); Mpio. 1.5 long, ca. 2 mm wide, dehiscing by longitu­ de Yarumal, Vereda EI Colgadero, Lozano C. dinal slits; staminode absent. Ovary ovoid, ca. 7 2172 (COL); Paramo de Sonson, Daniel 3444 mm long, ca. 4 mm wide, densely pilose, style (US); Daniel 3445 (US).-Cauca: Cordillera ca. 2 cm long, stigma stomatomorphic. Fruit a Central, eastern slope, Moscopan, valley of Rio fleshy capsule, pendent when ripe, ovoid when San Jose, Aguabonita, Cuatrecasas 23465 (F, immature, loculicidally dehiscent, becoming bi­ US); Cordillera Central, western slope, city of valved, valves reflexed, reaching a 1800 angle Rio Palo, Quebrada Santo Domingo, Cuatreca­ when mature, ca. 2 cm long, ca. 1.5 cm wide, sas 19304 (A, F, NY, US); Paramo de Las Pam­ pilose. Seeds numerous, fusiform, ca. 1 mm pas, between El Boquer6n and La Hoyola, [d­ long, ca. 0.3 mm wide, longitudinally striate, robo et aZ. 3837 (COL); Paramo de Las Pampas, dark purple. between Letreros and Santo Domingo, ldrobo et aZ. 3376 (COL, P); Mpio. de Coconuco, Valle PHENOLOGY: Collected in flower throughout de Paletara, Parque Natural Nacional de Purace, the year; in fruit in January, April, July, October, Lozano C. 3492 (COL); road Popayan-San and December. Agustin, km 70, Parque Nacional Purace, Lon­ DISTRIBUTION: Colombia, Ecuador, Venezuela, dono & Kvist 90 (AAU, COL, QCA, US).­ and northeru Peru; Andean slopes in wet mon­ Cundinamarca: below Saito de Tequendama, SE tane forests; 800-3500 m (FIGURE 2). of Santandercito, Uribe U. 1016 (COL, US); CLARK: ALLOPLECTUS MONOGRAPH 191

Cordillera Oriental, San Miguel, S of Sibate on docarpus, E of Nudo de Cajanuma, B¢gh 86568 road to Fusagasuga, Ian 35-36, Barclay et al. (AAU); Parque Nacional Podocarpus, S of Loja, 3395 (NA, US); paramos E of Guasca, toward E of Nudo de Cajanuma, (2)llgaard et al. 58010 Gacheta, 50 Ian NW of Bogota, Ariste-Joseph (AAU, MO, NY); Parque Nacional Podocarpus; s.n. (US); forest above Fusagasuga, Tracey 329 Nudo de Cajanuma, Yanez 91 (QCA, QCNE); (K); forest SE of Santandercito, near El Ermi­ Podocarpus National Park, area around Lagunas tano, Uribe U. 4915 (COL); forest S of Santan­ Compadres, Pedersen et al. 104445 (LOJA); dercito, Uribe U. 2961 (COL); mountain range Saraguro, Huashapamba, Proafio 3733 (LOJA). S of Santandercito, Lagoon of Catamica, Uribe -Napo: Cant6n Quijos, Parroq. Cuyuja, be­ U. 3195 (US); mountains below Tequendama, tween the town of Cuyuja and the confluence of between Cerro El Ermitafio and La Vencedora, Rio Tablon and Rio Quijos, northern border of Uribe U. 5568 (COL); Fusagasuga, Triana 607 Reserva Ecol6gica Antisana, Clark et al. 5625 (US); Paramo de San Miguel, western Andean (QCA, QCNE, US); road Papallacta-Baeza, be­ slopes, Duque J. 3454 (COL, COL); San Mi­ tween Cuyuja and Papallacta, Palacios 2573 guel, Schneider 686 (S); Santandercito, La Ram­ (MO, QCNE); Jaramillo 5314 (GB, MO, QCA); bla, Silva 272 (COL); Santandercito, La Rambla, van der Werif & Palacios 9620 (AAU, MO, Schneider 784 (COL); Sibate, Pennell 2496 QCNE, US); (2)llgaard & Eriksen 90304 (AAU, (NY); Tequendama, Pennell 2647 (GH, NY); QCA, QCNE); Palacios & van der Werff 3678 Tequendama, Fusagasuga, San Fortunato, Triana (AAU, MO, QCNE, US).-Thnguragua: Spruce 4163 (BM); Villeta to Utica highway, Garda 5179 (BM, C, E, G, GH, GOET, K, NY, OXF, Barriga 17203 (US); Triana 2499 (P).-Huila: P, W-2 sheets); Comunidad Viscaya; 7-11 Ian Rio Balsillas, Rusby & Pennell 786 (GH, NY); SW of Banos, Clark et al. 7744 (AAU, CAS, E, Cordillera oriental, E of Neiva, Rusby & Pennell F, K, MO, NY, QCA, QCNE, SEL, US, W); Rio 893 (NY); Cordillera Oriental, ridge NW of Vascun Valley; northern slopes of Volcan Tun­ Quebrada la Candela, Rio Naranjo, 20 kIn SW gurahua, Clark et al. 7720 (AAU, BM, E, F, of San Agustin, Fosberg 20121 (US); Mpio. de MO, NY, QCA, QCNE, SEL, US, WU); Clark Gigante, between Vereda Ventanas and Paramo et al. 7726 (QCNE, US); vicinity of Patate, Ha­ de Miraflores, Fernandez A. et al. 14870 (COL); cienda Leito, Asplund 8072 (S).-Zamora-Chin­ Fernandez A. et al. 14881 (COL).-Magdalena: chipe: road Loja-Zamora, Ian 12-14, Dodson & San Sabastilin de Rabago, Romero C. 937 (F, Thien 1382 (US, WIS). Peru-Piura: Provo US).-Meta: Sumapaz, western Andean slopes, Huancabamba, Los Llanos-Chorro Blanco (Sa­ Quebrada el Buque, Dlaz P. et al. 2826 (MO).­ palache-Chiguelas), Sagastegui et al. 16774 Narino: Mpio. Pasto, 3 Ian E of Dolores, Ra­ (US). Venezuela-Aragua: Parque Nacional Pit­ mirez P. & Cuayal M. 3954 (NY); Mpio. Pasto, tier, Fila de Paraiso, above Portuchuel0, Steyer­ Correg. El Encano, Ian 28-33 via Sibundoy, Ra­ mark 91523 (US, VEN).-Falcon: Sierra de San mIrez P. 5365 (US); Mpio. Pasto, Correg. Santa Luis, near the Parador Hotel, S of La Tabla, Barbara, headwaters of Rio Bobo, Ramirez P. & Steyermark 98977 (G, VEN); Steyermark 98948 Cuayal M. 4892 (US).-Quindio: El Roble, Tri­ (K, NY, P, VEN).-Tachira: Dist. Junin, between ana 4163 (BM); San Juan-Quindio, Andre 2045 EI Mirador and Potrero Buena Vista, Ruiz T. & (K, NY).-Risaralda: Cordillera Central, Mpio. Lopez F. 8912 (MERF, US); Quebrada de Pal­ de Santa Rosa, western Andean slopes, between mar, base of Paramo de Tama, 2 Ian above Be­ Hacienda La Sierra and Termales de Santa Rosa, tania and 7 Ian above Villapaez, Steyermark Jaramillo M. et al. 6026 (COL).-Tolima: Cor­ 57247 (F, US); slopes below paramo from Tama, dillera Central, Mpio. de Ronces Valles, Paramo near Colombian-Venezuelan border, Steyermark de Yerbabuena, Camargo G. 7437 (COL).-Val­ et al. 98427 (MO, NY, US, VEN).-Yaracuy: Ie del Cauca: El Silencio, Yanaconas, Garcia Quebrada Amparo, El Amparo, near Candelaria, 33806 (COL, US); Mpio. de Riofrio, Correg. de 7-9 Ian N of Salom, Steyermark & Espinosa Fenicia, El Roblal, Paramo del Duende, Vargas 111158 (SEL, VEN); Cerro La Chapa, N of Nir­ 3980 (COL). Ecuador-Azuay: Eastern Cordil­ gua, Meier 4197 (US); Steyermark & Bunting lera, between Ona and Rio Yacuambi, Prieto 97648 (COL, US, VEN). 212 (AAU, NY, S, US); road Sigsig-Gualaquiza, Alloplectus hispidus is distinguished from Harling et al. 8204 (GB, QCA, SEL).-Loja: other congeners by a prolifically branched Poortmann 143 (P-2 sheets); road to antennas climbing habit. Some populations have been ob­ from Lorna del Oro, between San Lucas and served to cover the entire base of a tree trunk to Saraguro, Pedersen & Lozano 104349 (LOJA); 5 m (J.L. Clark et al. 7720). The yellow flowers Saraguro-Loja, Ian 40.2, J¢rgensen et al. 1374 are similar in shape to the red flowers of A. in­ (LOJA, SEL, US); Parque Nacional Podocarpus, flatus, but the pouched region is not as pro­ Cajanuma, path towards Lagunas del Compadre, nounced (FIGURE 3B). The valvate to connivent Tirado 1529 (QCNE, US); Parque Nacional Po- calyx lobes of A. hispidus are useful in distin- 192 SELBYANA Volume 25(2) 2005

guishing it from the spreading calyx lob~s of .A. Seeds numerous, fusiform, 1 mm long, 0.3 mm aquatilis. These two species are allopatnc wIth wide, longitudinally striate, dark brown. the latter being restricted to the eastern flanks of PHENOLOGY: Flowering from February the Venezuelan Andes. through December; fruiting from August Alloplectus inflatus J.L.Clark & L.E.Skog, No­ through November. von 15(1): 72. 2005. TYPE: Colombia­ DISTRIBUTION: Alloplectus infiatus is known Cundinamarca: Muchindote Valley, east from the Andes of Colombia. It has been col­ side of Quebrada Negra, 13 km NE of lected in the Cordillera Central (Depts. Caldas, Gacheta, 2800 m, 4 Jul 1944, Grant 9596 Cauca, and Tolima) and the Cordillera Oriental (holotype: COL!; isotypes: MO! NA-2 (Depts. Boyaca, Cundinamarca, and Meta) at sheets! NY! US! WIS). FIGURE 4. 2000-3500 m (FIGURE 2). Shrubs terrestrial or epiphytic, stems scan­ REPRESENTATIVE SPECIMENS: Colombia-Boy­ dent, sparingly branched; to 2 m tall, to 0.8 cm aca: Cordillera Oriental, NE of Arcabuco, divid­ in diameter, woody, internodes 1-5 cm long, te­ ing line between Boyaca and Santander del Sur, rete, densely hirsute, leaf scars slightly raised Garda Barriga & Jaramillo M. 20261 (F, GH, from the stem surface, nectary glands at base of US).-Caldas: Cordillera Central, Alaska, above petiole present. Leaves opposite, usually eq~al Salento, Pennell 9383 (GH, PH); western slopes in a pair; petioles 0.5-2 cm long, densely ~Ir­ of the Cordillera Central, valley of Rio Otun, sute; blades coriaceous when dry, broadly ellIp­ Pefia Bonita, Cuatrecasas 23333 (F, US).-Cau­ tic, 3-10 cm long, 1-5 cm wide, base somewh~t ca: Eastern slopes of the Cordillera Central, re­ rounded to acute, apex obtuse to acute, margm gion of Moscopan, valley of Rio San Jose, serrate, adaxially dark green, tomentose, abaxi­ Aguabonita, Cuatrecasas 23484 (F, US); road ally light green to red, sparsely to densely hispid Popayan-La Plata, km 100, Researva Forestal de especially on the veins, lateral v~ins 3-5 per Fauna y Flora Merenberg, Murcia 12 (COL).­ side. Flowers solitary, non-resupmate, bracte­ Cundinamarca: San Miguel, near Sibate, Hawkes oles absent; pedicels longer than the petiole, 2- & Garcia Barriga 23 (K, US); between Pueb­ 7 cm long, pilose, pedicel glands absent, flower loviejo and Hoerias, near Boca de Monte, Uribe posture pendent at anthesis. Calyx lobes 5, near­ U. 6387 (COL); Cerros de San Miguel, SW of ly free, basally appressed to adjacent lobe, erect, La Sabana de Bogota, Uribe U. 2594 (COL, subequal, broadly ovate, 15-20 mm long, 5-~0 US); Cordillera Oriental, Moquentiva valley, 14 mm wide, base truncate, apex attenuate, margm km NW of Gacheta, Grant 9522 (NA, US); laciniate-serrate, red, outside densely pilose, in­ Mpio. Sibate, EI Pefi6n, Hertas & Camargo side sparsely pilose. Corolla 4.0-5.5 cm long, 5007 (COL); Paramo de Guasca, Schneider tubular, tube ca. 4 cm long, gibbous basally on 1196 (COL); vicinity of San Miguel, Weaver & upper surface, spur absent, long axis o! co.rolla Kennedy 1567 (GH).-Meta: eastern slopes of oblique relative to calyx; base ca. 1 cm m d~am., the Cordillera Central, Sunapas, quebrada El middle ampliate, becoming apically ventncose Buque, Dfaz P. 2677 (MO); main tributary of on lower surface, throat constricted, not appear­ Rio Grande, S of Cordillera de Las Cruces, ing laterally compressed, ca. 1 cm wide at southern slopes of Paramo de Sumapaz, Fosberg mouth, outside densely pilose, inside mostly gla­ 20862 (US); Mpio. Quetame, Guayabetal police brous, with glandular trichomes at throat, inte­ inspection station, road to Calvario, Lozano C. rior spotting absent, limb reflexed, subregular, 4063 (COL), Pefiuela & Clavijo 16 (COL).­ ca. 1 cm wide, red, lobes nearly equal, rotund, Tolima: near Quindio highway, along divide, spreading, ca. 2 mm long, ca. 3.5 mm wide, Killip & Varela 34612 (COL, US). rounded, entire. Nectary gland bilobed, on ven­ Alloplectus infiatus resembles A. hispidus, but tral surface of ovary, glabrous. Filaments coil­ differs by the more inflated pouched region near ing after anthesis, ca. 1.5 cm long (free portion), the apex of the ~orolla. Both A. hispidus and A. adnate to base of corolla tube for ca. 2 mm, con­ infiatus are scandent epiphytes, but A. hispid'!s nate for 1 mm, forming an open sheath, gla­ can cover an entire tree trunk for 3-5 m WIth Its brous; anthers broader than long, 1 mm long, 2 prolifically branched stems in contrast to the mm wide, dehiscing by longitudinal slits; stam­ sparingly branched stems of A. infiatus. Other inode absent. Ovary ovoid, ca. 7 mm long, ca. characters that are useful for distinguishing A. 4 mm wide, densely pilose, style 1.4 cm long, infiatus are the dark red flowers and nearly :al­ glabrous, stigma shallowly bilobed. Fruit a vate calyx lobes with basally appressed margms. fleshy capsule, pendent when ripe, ovoi~ whe.n immature, 10culicidal1y dehiscent, becommg bI­ Alloplectus tessmannii Mansf., Repert. Spec. valved, valves reflexed, reaching a 1800 angle Nov. Regni Veg. 41: 149. 1936. TYPE: when mature, 1.5 cm long, 1.5 cm wide, pilose. Peru-Loreto: upper Rfo Marafion, Puerto CLARK: ALLOPLECTUS MONOGRAPH 193

1cm

1cm

FIGURE 4. Alloplectus inflatus J.L.Clark & L.E.Skog. A. Habit. B. Adaxial leaf pubescence. C. Corolla. D. Corolla opened to show stamens. E. Calyx. F. Calyx opened and corolla removed to show bilobed nectary gland. G. Immature fruit. H. Seeds. (A, B from Uribe 2594 (US); C-F from Huertas & Camargo 5007 (COL); G, H from Pennell 9383 (PH)). lllustration from Clark et al. (2005: fig. 2) reprinted with permission. 194 SELBYANA Volume 25(2) 2005

Melendez, below Pongo de Manseriche, REPRESENTATIVE SPECIMENS: Peru-Huanuco: 155 m, 18 Dec 1924, G. Tessrnann 1800 Dist. Chinchao, San Pedro de Carpish, near Car­ (ho10type: B, destroyed). TYPE: Peru­ pish tunnel, ca. 47 kIn N of Huanuco on the road Amazonas: Prov. Bongara, "Shillac," em­ to Tingo Maria, trail towards Hacienda Patti, bankment of Rio Utcubamba and Quebrada Clark et al. 8210 (SEL, US, USM).-San Mar­ Alba, ca. 10 kIn from the Korontachaca tin: Prov. Rioja, Dist. Naranjillo, sector San bridge along the old road to Herradura de Agustin, Sanchez V. & Dillon 8653 (F, US); Chachapoyas and Bagua, 2270-2490 m, Provo Rioja, Dist. Nuevo Cajamarca, Sanchez V. 12-13 Sep 1983, M. Leo L. 309 (neotype: & Dillon 8477 (F, US). MO!; isoneotype: FLAS!). Alloplectus tessrnannii is infrequently collect­ Subshrubs epiphytic, stems scandent, rarely ed and specimens of this species in flower are branched; to 1 m long, 4-6 mm in diam., sub­ rare. The pouched region of the corolla is medial woody, internodes 2-5 cm long, terete, with in contrast to the nearly terminal pouch of other dense villous pubescence, leaf scars slightly congeners. The stems of A. tessmannii are raised from the stem surface, nectary glands at sparsely branched and erect in contrast to the base of petiole present. Leaves opposite, nearly scandent stems of other congeners. It is sympat­ equal in a pair; petioles 1.5-5.0 cm long, villous; ric with A. hispidus, from which it is easily dif­ blades coriaceous when dry, ovate to oblong, ferentiated by the red flowers and sparsely 10-12 cm long, 4-5 cm wide, base acute, apex branched erect stems. acute, margin serrate, adaxially dark green, pi­ Alloplectus weirii (Kuntze) Wiehler, Phytologia lose, becoming more pilose on venation, abaxi­ 27: 327. 1973. Crantzia weirii (Kuntze) ally pale green, uniformly pilose, lateral veins Fritsch in Engler & Prantl, Nat. Pflanzen­ 6-8 per side. Flowers solitary, bracteoles absent; fam. 4(3b): 168. 1894. Columnea weirii pedicels shorter than the petiole, to 1 cm long, Kuntze, Revis. Gen. PI. 2: 471. 1891. TYPE: densely pilose, pedicel glands absent, flower Colombia-Cundinamarca: woods near Pa­ posture horizontal at anthesis. Calyx lobes 5, cho, 1888, Weir 217 (holotype: K!). nearly free, base of each lobe appressed to ad­ FIGURE 5. jacent lobe, with the margin curved inward, erect, nearly equal, broadly oblong, to 2.5 cm Alloplectus lindenii (Kuntze) Wiehler, Phytolo­ long, to 1 cm wide, base truncate, apex acute, gia 27: 327. 1973. Columnea lindenii margin laciniate, green, outside densely villous, Kuntze, nom. illeg. [non Columnea lindenii inside villous. Corolla to 3 cm long, tubular, Brong. ex Lem.], Revis. Gen. PI. 2: 471. tube 2.6 cm long, gibbous basally on upper sur­ 1891. Crantzia lindenii (Kuntze) Fritsch in face, spur absent, long axis of corolla oblique Engler & Prantl, Nat. Pflanzenfam. 4(3b): relative to calyx; base ca. 6 mm in diam., middle 168. 1894. TYPE: Colombia-Provincia de ampliate and curved, becoming ventricose on Soto [Dept. Santander], 1800-2100 m, Dec lower surface, throat slightly constricted, not ap­ 1853, L. Schlim 1089 (lectotype, here des­ pearing laterally compressed, outside glabrous at ignated: K!; isolectotypes: G! PI). base, becoming densely pilose distally, inside Columnea schlimii Kuntze, Revis. Gen. PI. 2: glabrous, interior spotting absent, limb erect, zy­ 471. 1891, nom. nud. pro syn., non Col­ gomorphic, to 4 mm wide, uniformly red, lobes umnea schUmii (Planch. & Linden) Kuntze nearly equal, rotund, 1 mm long, 1.5 mm wide, (see discussion below). rounded, entire. Nectary gland bilobed, on ven­ tral surface of ovary, glabrous. Filaments coil­ Vines epiphytic, stems climbing or sprawling, ing after anthesis, 1 cm long (free portion), ad­ adhering to other plants by adventitious roots, nate to base of corolla tube for ca. 4 mm' con­ frequently branched; growing 2-5 m above nate for 1 mm, forming an open sheath, gla­ ground, stems 3-7 mm in diam., subwoody, in­ brous; anthers broader than long, 2 mm long, 2.5 ternodes 2-5 cm long, terete, glabrescent below, mm wide, dehiscing by longitudinal slits; stam­ dense yellowish-green pilose pubescence above, inode absent. Ovary ovoid, ca. 8 mm long, ca. leaf scars flush with stem surface, nectary glands 5 mm wide, densely pilose, style 2 cm long, pu­ at base of petiole present. Leaves opposite, equal bescent, stigma stomatomorphic. Fruit and or subequal in a pair; petioles 1-4 cm long, seeds not observed. densely pilose; blades subcoriaceous when dry, usually elliptic, sometimes slightly ovate, 5-11 PHENOLOGY: Flowering in November. cm long, 2-4.5 cm wide, base acute to obtuse, DISTRIBUTION: Alloplectus tessmannii is apex acute, margin serrate-crenate, adaxially known from montane rain forests in Peru (Huan­ dark green, sparingly strigose, abaxially pale uco, Amazonas, San Martin, and Loreto) 1200- green, sometimes red, densely pilose, lateral 2400 m (FIGURE 2). veins 6-8 per side. Inflorescence not peduncu- CLARK: ALLOPLECTUS MONOGRAPH 195

AI,Wvlectus weirii (Kuntze) Wiehler. A. Habit. B. Lateral view of flower. C. Lateral view of immature fruit persistent calyx. Photographic images of field collection J.L. Clark & L. Bohs 8914 (US). late, flowers usually solitary, rarely 2 flowers per tund, 2 mm long, 2.5 mm wide, rounded, entire. node, bracteol~s lanceolate, 6 mm long, 0.2 mm Nectary gland on dorsal surface, usually single­ wide; pedicels Ishorter than the petiole, 0.5-1 cm lobed, rarely bilobed, glabrous. Filaments coil­ long, densely i pilose, pedicel glands absent, ing after anthesis, 1 cm long (free portion), ad­ flower posture ipendent at anthesis. Calyx lobes nate to base of corolla tube for 6 mm, connate 5, connate brit#IY at base, each appressed to ad­ for I mm, forming an open sheath, glabrous; an­ jacent lobe, fo~ded lengthwise with the margin thers broader than long, 2 mm long, 3 mm wide, curved inwardj erect, 4 equal, broadly ovate, to dehiscing by longitudinal slits; staminode ab­ 2.0 cm long, t~ 1.5 cm wide, base truncate, apex sent. Ovary ovoid, ca. 7 mm long, ca. 6 mm obtuse to acut~, margin fimbriate, the teeth to wide, pilose, style 1.5 cm long, sparingly pilose, 1.0 cm long and ca. 0.2 cm wide, all green to stigma stomatomorphic. Fruit a fleshy capsule, all red, outsideidensely pilose, inside densely pi­ pendent, ovoid when immature, loculicidally de­ lose; fifth (ve~tral) lobe smaller and narrower, hiscent, becoming bivalved when mature, 2 cm broadly ovate,! to 2 cm long, to 0.5 Icm wide. long, 1.5 cm wide, pilose. Seeds numerous, el­ Corolla to 2.5! cm long, tubular, tube to 2 cm lipsoid, I mm long, 0.3 mm wide, longitudinally long, gibbous ~asally on upper surface, spur ab­ striate, light red. sent, long axis iof corolla oblique relative to ca­ PHENOLOGY: Collected in flower year round lyx; base ca. mm in diam., middle ampliate .5 except for the months of October and December; and ventricose Ion the lower surface, throat con­ collected in fruit during the months of January, stricted, not appearing laterally compressed, 6- March, April, June, July, October, November, 8 mm wide (at mouth), outside densely villous, and December. inside glabrous, interior spotting absent, limb spreading, zygomorphic, to 9 mm wide, pinkish­ DISTRIBUTION: Alloplectus weirii is known yellow to orangish-red, lobes nearly equal, ro- from the Andes of NW South America (Bolivia, 196 SELBYANA Volume 25(2) 2005

Peru, Ecuador, and Colombia) and Costa Rica at to Las Lagunas de Sardina Yacu, between Rios 500-2500 m (FIGURE 2). Sardina and Volcan, Ceron M. & Montalvo 32721 (QAP); Canton Morona, Cordillera del REPRESENTATIVE SPECIMENS: Costa Rica-AI­ Cutucu, Asociacion Shuar Sevilla, Comunidad ajuela: 17-20 km NNW of San Ramon on road Angel Ruby, behind Cerro Ifiiak naint, Suin et towards San Lorenzo, 4-7 km N of Balsa, Lies­ al. 1805 (MO, QCNE).-Napo: 1 km E of oil ner 14694 (CR, US); Alajuela-San Ramon, RIO pump station on road Papallacta-Baeza, along Cataratas y Cataratitas, Gomez et al. 20789 (CR, Rio Papallacta, Holm-Nielson et al. 6969 (AAU, US); road San Ramon-Baja Rodriguez, km 19.5 NY, S); environs of Baeza, Besse et al. 1155 from San Ramon, Croat 78842 (MO); road San (SEL); Besse et al. 1161 (SEL); Canton Archi­ Ramon-Balsa, 16.7 km N of bridge over Que­ dona, Sumaco Napo-Galeras National Park, base brada Volio, 7.5 km N of bridge over Rio Balsa, of Volcan Sumaco, between Pacta Sumaco and Stevens 13913 (MO, US); N of San Ramon, ca. main road to Loreto, Clark 2307 (QCNE, US); 4 km N of Balsa along road to Colonia Palmar­ Baeza, Kvist et al. 60393 (AAU, US); Canton ena, Lellinger & White 1253 (SEL, US); Canton Quijos, at the Rio Cosanga near Cosanga, Kirk­ San Ramon, Dist. Angeles, Los Angeles, Colon­ bride & Chambra R. 4009 (AAU, Q, US); Can­ ia Palmarena, Cuenca media de Rio San Lor­ ton Quijos, Parroq. Cosanga, Yanayacu Biolog­ enzo, camino a la mina de yeso, Herrera 4924 ical Station and Center for Creative Studies, 5 (CR, INB, US); Rio Penas Blancas, near Refu­ km W of Cosanga, Clark & Greeney 5788 gio Aleman, Haber 11298 (lNG, MO); trail to (COL, MO, QCNE, US); Clark & Greeney 5792 Volcan Muerto, Araya et al. 264 (lNG, MO); (AAU, MO, NY, QCA, QCNE, US); Cerro An­ Monteverde Reserve, Atlantic slope, Rio Penas tisana, 1 mile NE of Borja, Grubb 1271 (K-2 Blancas valley, Haber 1007 (US); Haber 852 sheets, NY); Canton EI Chaco, Proyecto Hidroe­ (US); Haber 996 (US); Obando et al. 167 (INB, lectrico Coca, Punta ST3, Rio Quijos, 10 km S US); Reserva Forestal, San Lorenzo, San Ra­ of Reventador, Palacios 5936 (MO); Canton mon, Carvajal 166 (US); Carvajal 368 (US).­ Quijos, western slopes of the Cordillera de Gua­ Guanacaste: Canton de Liberia, Parque Nacional camayos, edge of Rio Cosanga, Palacios 6351 Guanacaste, Cordillera de Guanacaste, Nueva (MO); Vicinity of Baeza, Dwyer & Macbryde Zelandia, Estacion San Ramon, Sendero la Te­ 9581 (MO, US).-Pastaza: along Rio Pastaza, pescuintle, Garda 189 (CR, INB, US). Boliv­ between Rio Tapa and Rio Mera, Steyermark ia-La Paz: Yungas, Rusby 2424 (NY). Colom­ 54882 (F, US); Lugo S. 1140 (GB, SEL); Mera, bia-Dept. unknown: Mutis 2249 (MA).-An­ along Rio Pastaza, Lugo S. 1124 (GB, SEL).­ tioquia: Kalbreyer 1456 (K); Mpio. de Caldas; Tungurahua: Banos, hotel "EI Otro Lado," Rio finca La Oculta, Escobar & Uribe 3621 (US); Verde (tributary of Rio Pastaza), near El Pailon Rio Calderas, Daniel 4471 (US).-Boyaca: re­ del Diablo, Clark & Rea 8016 (QCNE, US); gion of Mt. Chapon, extreme western part of Cant6n Banos; Parroquia Rio Negro; Sectorlas Dept. Boyaca, NW Bogota, Lawrance 357 (K, Palmeras, between Rio Estancia and Rio Pasta­ U).-Caldas: Cordillera Central, Salento, Pen­ za, Ceron M. & Davila 38680 (QAP); Canton nell 9081 (NY, PH, US).-Caqueta: western Banos, Parroq. Rio Verde, Sector Machay, for­ slopes of the Cordillera Oriental, Quebrada del ested trail from Banos-Puyo road towards Cas­ Rio Hacha, Cajon de Pulido, Cuatrecasas 8752 cada de San Miguel via San Agustin, Clark et (COL, US).-Cundinamarca: Cordillera Orien­ al. 5701 (AAU, COL, MO, NY, QCA, QCNE, tal, Caraucha Range, E of Ibama, 13 km E of SRp, US).-Zamora-Chinchipe: W of Rio Val­ Yacopi, near the Boyaca border, Grant 9184 ladolid, above Valladolid, Steyermark 54699 (F, (COL, NA, WIS); Tena, Laguna de Pedro Palo, US); Parroquia Zumba, Finca de Sandy Le6n, Uribe U. 1758 (GH).-Quindio: Mpio. de Fi1an­ Forest near Rio Tarrangami, J.L. Clark et at., dia, Brehmen, Vargas W. 3224 (COL).-Santan­ 8914 (QCNE, US). Peru-Cajamarca: Cutervo, der: road to Pamplona, km 12, Alto de los Pa­ Parque Nacional de Cutervo, San Andres de Cu­ dres, Garda Barriga & Jaramillo M. 20545 tervo, Smith & Diaz 1655 (US); La Palma, 10 (COL, SEL, U, US); Correg. Virolin, road Can­ km NW of Chirinos, Gentry et al. 61191 averales-Olival, Rio Ruicito, Londono & Kvist (MO).-Cutervo: San Andres de Cutervo, road 809 (US).-Tolima: Mariquita, La Palmilla, San Andres-Santo Tomas, D{az & Beltran 3341 Mariquita, Palmilla, Triana 4163 (BM); Alto de (US, USM); San Ignacio, Ruta San Ignacio-EI la Palmilla, Triana 2484 (BM, P, W).-Valle del Chaupe, Leiva et al. 1574 (F, NY, US); San Ig­ Cauca: Mpio. Restrepo, vereda de Roman, road nacio, San Jose de Lourdes, Santo Tomas, Cam­ Pavas-Restrepo, Ramos et al. 2574 (US); La pos & Corrales 3441 (US).-Cuzco: Conven­ Cumbre, Cordillera Occidental, Killip 5926 cion, Choquellohuanca, Marin 2116 (F, US).­ (NY); Pennell & Killip 5782 (NY, PH, US). Ec­ Loreto: Provo Coronel Portillo, Dtto. Padre uador-Morona-Santiago: Canton Macas, trail Abad, Rio Chino, W of restaurant Acapulco, CLARK: ALLOPLECTUS MONOGRAPH 197

Schunke V. 9143 (GH, MO, MO, NY, US).­ Peru-Loreto, upper Rio Maraii6n, delta of San Martin: Prov. Rioja, road Pedro Ruiz-Moy­ Rio Santiago, tierra firma forest, 1 Oct obamba, km 390, Venceremos, Smith & Vasquez 1924, G. Tessmann 4182 (B, no longer ex­ S. 4723 (K, US). tant). TYPE: Ecuador-Morona-Santiago, While many other taxonomists working in Cordillera Morire, San Miguel de los Cu­ Gesneriaceae did not avoid homonyms (e.g., yes, 2400-3400 m, 7 Jan 2001, J.L. Clark Linden), Kuntze (1891) recognized the potential & C. Morocho 5847 (neotype, here desig­ problems and attempted to avoid them. The nated: US!; isoneotypes: HA! QCNE!). name "Columnea schlimii Kuntze" appeared in =Glossoloma altescandens (Mansf.) J.L. the text (Kuntze 1891) of his description of Col­ Clark, comb. nov. umnea lindenii Kuntze. Thus, these two names Alloplectus ambiguus Urb., Symb. Antill. 1: 408. are of the same species. It is likely that Kuntze 1899. =Columnea ambigua (Urb.) B.D. mentioned the name Columnea schlimii in the Morley, Proc. Roy. Irish Acad. 74B: 412, text, because he had already annotated collec­ 423. 1974. tions with this name (e.g., L. Schlim 1089 from Alloplectus ambiguus var. chlorosepalus Urb., K!). It is presumed therefore that he cited the Symb. Antill. 1: 408. 1899. =Columnea name to avoid any confusion. Realizing that this ambigua (Urb.) B.D.Morley, Proc. Roy. would be an undesirable homonym, Kuntze in­ Irish Acad. 74B: 412, 423. 1974. cluded "Columnea schlimii Kuntze" in the text Alloplectus ambiguus var. erythrosepalus Urb., of his description and proposed the name Col­ Symb. Antill. 1: 408. 1899. =Columnea umnea lindenii Kuntze. ambigua (Urb.) B.D.Morley, Proc. Roy. Alloplectus weirii is distinguished from other Irish Acad. 74B: 412, 423. 1974. members of the genus by deeply fimbriate calyx Alloplectus ambonensis L.E.Skog, Ann. Missou­ margins (FIGURE 5C). In the absence of mature ri Bot. Gard. 65: 795. 1978 [1979]. TYPE: flowers, A. weirii is easily confused with many Panama-Border of ChiriqUI and Bocas del of the Columnea species from section Ortholo­ Toro, road along Cerro Colorado ridge, J.P. ma (e.g., Columnea minor) because of the fim­ Folsom, G. Smail, and R. Robbins 4774 briate calyx margins and isophyllous opposite (holotype: US!; isotypes: MO-2 sheets! leaves. The corolla is anomalous in the genus US!). =Drymonia ambonensis (L.E.Skog) because of its nearly urceolate shape and lack of J.L.Clark, comb. nov. a well-defined pouched region (FIGURE 5B). Alloplectus angustifolius DC., Prodr. 7: 546, 1839. Columnea angustifolia (DC.) Kuntze, Revis. Gen. PI. 2: 472. 1891. =Nematan­ AN ALLOPLECTUS NOMENCLATOR thus lanceolatus (Poir.) Chautems, Diss. Bot. 112: 189. 1988. Excluded or Uncertain Species Alloplectus aureonitens (Hook.) Stearn, Bull. The following is a list of published names of Brit. Mus. (Nat. Hist.) Bot. 4(5): 189. 1969. Alloplectus and their present taxonomic place­ Columnea aureo-nitens Hook., Bot. Mag. ment. An equal sign (=) precedes the currently 73: pI. 4294. 1847. Collandra aureonitens accepted name (bold type). Combinations made (Hook.) Hanst., Linnaea 26: 209. 1854. by previous authors are noted with place of pub­ Dalbergaria aureonitens (Hook.) Wiehler, lication. Also noted (e.g., "fide") are heterotyp­ Phytologia 27: 316. 1973. TYPE: Plate 4294 ic synonyms recognized by previous authors. in J.W Hooker, Bot. Mag. 73. The "native specimens are in our Herbarium from Co­ Alloplectus affinis (C.V.Morton) Stearn, Bull. lumbia" as mentioned by Hooker in the text Brit. Mus. (Nat. Hist.) Bot. 4(5): 189. 1969. were not seen by Leeuwenberg (1958), nor Columnea affinis C.Y.Morton, Fieldiana, was I able to locate a specimen during a Bot. 28: 529. 1953. TYPE: Venezuela-Ar­ 2002 visit to the Royal Botanic Gardens, agua: El Portachuelo, between Ocumare de Kew. The colored lithograph and text in la Costa and Maracay, H. Pittier 10130 Hooker's description allow this name to be (US I), nom. illeg., later homonym of Allo­ attributed as a synonym of Columnea san­ plectus affinis Mansf. =Columnea sangui­ guinea (Pers.) Hanst. This is a common spe­ nea (Pers.) Hanst., Linnaea 34: 384. 1865. cies throughout Colombia where the plant Alloplectus affinis Mansf., Repert. Spec. Nov. in cultivation at Kew was originally col­ Regni Veg. 41: 147. 1936. =Drymonia af­ lected. =Columnea sanguinea (Pers.) finis (Mansf.) Wiehler, Phytologia 27: 324. Hanst., Linnaea 34: 384. 1865. 1973. Alloplectus baguensis L.E.Skog, Opera Bot. 92: Alloplectus altescandens Mansf., Repert. Spec. 225. 1987. TYPE: Peru-Amazonas: Bagua, Nov. Regni Veg. 41: 148. 1936. TYPE: ca. 25 km (by trail) E of La Peca, along 198 SELBYANA Volume 25(2) 2005

stream, 1894 m, 13 Aug 1978, P. Barbour Club 4: 238. 1895. TYPE: Bolivia-La Paz: 2973 (holotype: US!; isotypes: C! MO!). Yungas, Oct 1890, Bang 515 (holotype: =Glossoloma baguense (L.E.Skog) J.L. NY!; isotypes: BM! E! F! GH-2 sheets! K! Clark, comb. nov. MANCH (photo: US!), MO! NY! PH! US- Alloplectus bicolor (Kunth) G.Don in Loudon, 2 sheets! W (photo: US!), WU (photo: Encycl. PI. new ed.: 1402. 1855. This name US 1); fragment: F!). =Glossoloma boli­ was published as a new combination for vianum (Britton ex Rusby) J.L.Clark, Besleria bicolor Kunth, but it has been cited comb. nov. as Alloplectus bicolor G.Donf. by Mabber­ Alloplectus bracteatus Linden, Etabl. Linden, ley (1981). =Glossoloma bicolor (Kunth) Prix-courant no. 5: 2. 1850 (see Stafieu & J.L.Clark. Cowan 1981: Vol. 3, entry 4620a). I was Alloplectus bicolor (Kunth) Klotzsch ex Hanst., not able to locate herbarium specimens with Linnaea 34: 377. 1865. Besleria bicolor this name attributed to Linden during a Kunth in H.B.K., Nov. Gen. Sp. PI. 2: 398 2002 visit to the National Botanic Garden (folio), 319 (quarto). 1817 [1824]. Alloplec­ of Belgium (BR), and the publication where tus kunthii G.Don, Gen. Hist. 4: 655. 1837 this name appears is not known to exist [1838]. Columnea bicolor (Kunth) Kuntze, (Stafleu & Cowan 1981). The names Allo­ Revis. Gen. PI. 2: 472. 1891. Crantzia bi­ plectus bracteatus Linden and Centrosolen­ color (Kunth) Fritsch, Bot. Jamb. Syst. 29 ia bracteatus Planch. are probably the same (Beibl. 65): 9. 1900. TYPE: Colombia-Cau­ species. It is assumed here that these names ca/Narino: Villa La Erre, plains of Sacan­ are of the same species advertised in Lin­ donoy between town of Almaguer and Pas­ den's seed catalogs. =Nautilocalyx brac­ to, 1829-2195 m, Oct 1801, A. Bonpland teatus (Planch.) Sprague, Bull. Misc. In­ 2118 (holotype: P! holotype fragment: F!). form. 1912: 88. 1912. =Glossoloma bicolor (Kunth) J.L.Clark, Alloplectus calcaratus (Kunth) G.Don, Gen. comb. nov. Hist. 4: 655, 1837 [1838]. Besleria calcar­ Alloplectus bicolor Linden, Suppl. Extrait Cat. ata Kunth in H.B.K., Nov. Gen. Sp. Pl. 2: Gener. No. 83 et 24. 1869 (see Stafieu & 320 (folio), 399 (quarto). 1817 [1824]. Cowan 1981: Vol. 3, entry 4626). There are =Gasteranthus calcaratus (Kunth) Wieh­ no cited specimens in the seed catalog ler subsp. calcaratus, Selbyana 1(2): 154. where this name was published, and I was 1975. not able to locate annotated herbarium spec­ Alloplectus calochlamys Donn.Sm., Bot. Gaz. imens with this name during a 2002 visit to 27: 437. 1899. Columnea calochlamys the National Botanic Garden of Belgium (Donn. Sm.) C.V.Morton, Bol. Soc. Venez. (BR). A vague description cites the similar­ Cienc. Nat. 23: 78: 1962. TYPE: Guatema­ ities of Linden's A. bicolor with two species la-Alta Verapaz: Sacoyoju, often on lime­ that are currently recognized as members of stone in open pJaces or thickets, 1090 m, the genus Corytoplectus (Alloplectus schli­ May 1879, H. von Tuerckheim 456 (lecto­ mii Planch. & Linden and Alloplectus spe­ type: US! designated by Gibson 1974). ciosus Poepp.). Linden's generic concept of Smith (1899) cited collections from two dif­ Alloplectus included what currently is rec­ ferent localities with the same number (H. ognized as Corytoplectus, and, in the ab­ von Tuerckheim 456). The collections from sence of a detailed description and lack of Pansamahi were syntypes (GH! K! NY! P! locality information, it is difficult to assign W! US!), until Gibson (1974) chose the col­ this name to a species. This name is most lection from Sacoyoju as the lectotype. likely a member of the congeners men­ Cobananthus Wiehler is in a clade that is tioned in Linden's vague 1869 description. well supported and separate from Alloplec­ =Corytoplectus sp. tus. It is more closely related to the primar­ AZloplectus bicolor Poepp. in sched. in Hanstein, ily Central American genera Alsobia Hanst. Linnaea 34: 370. 1865, nom. nud. pro syn. and Episcia Mart. than to either Columnea Hanstein cited Poeppig's name in reference or Alloplectus (Clark & Zimmer 2003; to it being seen on an herbarium collection. Clark et al. in press). The fruits of Coban­ Hanstein included this name as a synonym anthus are bivalved capsules with thick of Alloplectus speciosus Poepp. =Cory to­ non-refiexing carpel walls, whereas the plectus speciosus (Poepp.) Wiehler, Phy­ fruits of most other Episcieae genera (e.g., tologia 27: 313. 1973. Drymonia and Alloplectus) are fleshy cap­ Alloplectus bolivianus (Britton ex Rusby) Wieh­ sules with spreading refiexed carpel walls. ler, Phytologia 27: 327.1973. Columnea bo­ Other characters that distinguish this genus liviana Britton ex Rusby, Mem. Torrey Bot. are the actinomorphic, star-shaped, bright CLARK: ALLOPLECTUS MONOGRAPH 199

orange calyx, and the narrow tubular corolla maculatus Leeuwenb., Acta Bot. Need. 7: (cf., Wiehler 1977). =Cobananthus caloch­ 300. 1958. =Drymonia coccinea (Aubl.) lamys (Donn.sm.) Wiehler, Selby ana 2: Wiehler, Phytologia 27: 324. 1973. 94, pI. 28B. 1977. Alloplectus concolor Hook., Bot. Mag. 74: pI. Alloplectus calotrichus (Donn.Sm.) Stearn, Bull. 4371. 1848. TYPE. Plate 4371 in W.J. Hook­ Brit. Mus. (Nat. Hist.) Bot. 4(5): 189. 1969. er, Bot. Mag. 74. 1948. Hooker wrote, "we Ortholoma calotrichum (Donn.Sm.) Wieh­ presume, an inhabitant of Brazil, having ler, Phytologia 27: 321. 1973. Trichantha been sent to Kew by Mr. Galeotti in 1846 calotricha (Donn.Sm.) Wiehler, Selbyana ... without any indication of its locality." 1(1): 34. =Columnea calotricha Donn.Sm., It is not clear why Hooker thought that this Bot. Gaz. 40: 9. 1905. species is from Brazil, because Henri G. Alloplectus capitatus Hook., Bot. Mag. 75: pI. Galeotti (1841-1858) only collected in 4452. 1849. Columnea capitata (Hook.) Cuba and Mexico. Perhaps, Hooker meant Kuntze, Revis. Gen. PI. 2: 472. 1891. George Gardner (1812-1849), who had col­ Crantzia capitata (Hook.) Fritsch in Engler lected in Brazil during Hooker's tenure at & PrantI. Nat. Pfianzenfam. 4(3b): 168. Kew. Nevertheless, I was not able to locate 1894. =Corytoplectus capitatus (Hook.) a Galeotti or a Gardner herbarium specimen Wiehler, Phytologia 27: 313. 1973. of this species during a 2002 visit to the Alloplectus carpishensis J.L.Clark & LSalinas, Royal Botanic Gardens, Kew. =Nematan­ Novon 15: 70. 2005. TYPE: Peru-Huanuco: thus lanceolatus (Poir.) Chautems, Diss. Chinchao, San Pedro de Carpish, above Bot. 112: 189. 1988. Carpish tunnel, ca. 47 km N of Huanuco on Alloplectus congestiflorus (Donn.Sm.) C.Y. the road to Tingo Marfa, 09°43' 14"S, Morton, Field Mus. Nat. Hist., Bot. Ser. 18: 76°06'53"W, 2770-2900 m, 1 Nov 2001, 1. 1143. 1938. Besleria congestiflora Salinas 224 (holotype: USM; isotype: US!). Donn.Sm., Bot. Gaz. 61: 379. 1916. =Ru­ =Glossoloma carpishense (J.L.Clark & fodorsia congestiflora (Donn.Sm.) Wieh­ I.Salinas) J.L.Clark, comb. nov. ler, Selbyana 1(2): 146. 1975. Alloplectus chrysanthus Planch. & Linden, FI. Alloplectus congestus Decne. ex Linden, BtabI. Serres Jard. Eur. 8: 211. 1853. Columnea Linden, Prix-courant no. 4. 1849 (see Sta­ chrysantha (Planch. & Linden) Kuntze, Re­ fieu & Cowan 1981: Vol. 3, entry 4620a). vis. Gen. PI. 2: 472. 1891. No locality or There are no cited specimens in the seed type specimen designated. TYPE: Venezue­ catalog where this name was published, and la-Merida: dirt road towards Paramo de I was not able to locate annotated herbarium Mucumbis, between Torondoy and Pico specimens with this name attributed to De­ Aguila, near entrance to Parque Nacional caisne during a 2002 visit to the National Culata, 2800-2900 m, 16 Oct 2002, J.L. Botanic Garden of Belgium (BR). A vague Clark & S. Yustiz 6872 (neotype, here des­ description mentions that the corolla is pink ignated: US!; isoneotypes: MER! NY! and that the wild origin of this species is PORT! SELl VEN!). =Glossoloma chry­ Colombia. The corolla color and locality santhum (Planch. & Linden) J.L.Clark, agree with Corytoplectus congestus (Linden comb. nov. ex Hanst.) Wieh1er. Furthermore, it is not Alloplectus circinatus Mart., Nov. Gen. Sp. PI. clear if Hanstein differentiated Linden's 3: 56, pi. 223, Fig. 2. 1829. Columnea cir­ species from Decaisne's Alloplectus conges­ cinata (Mart.) Kuntze, Revis. Gen. PI. 2: tus. It is assumed here that these names are 472. 189l. Crantzia circinata (Mart.) of the same species advertised in Linden's Fritsch in Engler & Prantl, Nat. Pfianzen­ seed catalogs. =Corytoplectus congestus fam. 4(3b): 168. 1894. =Drymonia cocci­ (Linden ex Hanst.) Wiehler, Phytologia nea (Mart.) Wiehler, Phytologia 27: 324. 27: 313. 1973. 1973. Alloplectus congestus Linden ex Hanst., Linnaea Alloplectus coccineus (AubI.) Mart., Nov. Gen. 34: 37l. 1865. Columnea congesta (Linden Sp. PI. 3: 57 [text]. 1829. 3: 189 [index]. ex Hanst.) Kuntze, Revis. Gen. PI. 2: 472. 1832. Besleria coccinea AubI., Hist. PI. 189l. Crantzia congesta (Linden ex Hanst.) Guiane 2: 632, pI. 255. 1775. Lophalix coc­ Fritsch in Engler & Prand, Nat. Pfianzen­ cinea (AubI.) Raf., Sylva Tellur.: 70. 1838. fam. 4(3b): 168. 1894. =Corytoplecfus con­ Columnea coccinea (AubI.) Kuntze, Revis. gestus (Linden ex Hanst.) Wiehler, Phy­ Gen. PI. 2: 472. 1891. =Drymonia cocci­ tologia 27: 313. 1973. nea (Aubl.) Wiehler, Phytologia 27: 324. Alloplectus coriaceus (Oerst. ex Hanst.) Hanst., 1973. Linnaea 34: 363. 1865. Erythranthus cori­ Alloplectus coccineus (AubI.) Mart. var. fusco- ace us Gerst. ex Hanst., Linnaea 26: 183, 200 SELBYANA Volume 25(2) 2005

209, 215, 1854, Columnea coriacea (Oerst. Alloplectus cristatus (L.) Mart. var. brevicalyx ex Hanst.) Kuntze, Revis. Gen. PI. 2: 472. C.V.Morton, Contr. U.S. Natl. Herb. 29: 17. 1891. Crantzia coriacea (Oerst. ex Hanst.) 1944. =Crantzia cristata (L.) Scop., Intr. Fritsch in Engler & Prantl, Nat. Pflanzen­ Hist. Nat.: 173. 1777. fam. 4(3b): 168. 1894. A detailed descrip­ Alloplectus cristatus (L.) Mart. var. crenatus tion of this species name can be found in C.VMorton, Contr. U.S. Nati. Herb. 29: 17. Oersted (Centralamer. Gesner. 47, pI. 3, 1944. =Crantzia cristata (L.) Scop., Intr. 1858 [preprint]; 121, pI. 3, 1861 [reprint]). Hist. Nat.: 173. ]777. Oersted's species name and a copy of his Alloplectus cristatus (L.) Mart. var. epirotes illustration were published by Hanstein. Al­ Leewenb., Acta Bot. Need. 7: 298. 1958. though Hanstein attributed this name to TYPE: Guyana-Cuyuni-Mazaruni: Mt. Ay­ Oersted, it had not been published and is anganna, Pakaraima Mountains, 1000-1500 therefore treated as validated by Hanstein. m, 5-6 Feb 1955, B. Maguire, w.M.c. Bag­ =Drymonia coriacea (Oerst. ex Hanst.) shaw & c.K. Maguire 40594 (holotype: Wiehler, Phytologia 27: 325. 1973. NY!; isotypes: NY! U! US!). =Crantzia Alloplectus corymbosus Klotzsch ex Hanst., Lin­ epirotes (Leeuwenb.) J.L.Clark, stat. & naea 34: 371. 1865. Columnea corymbosa comb. nov. (Klotzsch ex Hanst.) Kuntze, Revis. Gen. Alloplectus cubensis (Urb.) Stearn, Bull. Brit. PI. 2: 472. 1891. Crantzia corymbosa Mus. (Nat. Hist.) Bot. 4(5): 189. 1969. Col­ (Klotzsch ex Hanst.) Fritsch in Engler & umnea sanguinea (Pers.) Hanst. var. cub­ Prand, Nat. Pflanzenfam. 4(3b): 168. 1894. ensis Urb., Symb. Antill. 2: 359. 1901. Col­ =Corytoplectus speciosus (Poepp.) Wieh­ umnea cubensis (Urb.) Britton, Torreya 5: ler, Phytologia 27: 313. 1973. 215. 1906. =Columnea sanguinea (Pers.) Alloplectus crenatilobus Manse Notizbl. Bot. Hanst., Linnaea 34: 384. 1865. Gart. Berlin 14(121): 38. 1938. TYPE: Ec­ Alloplectus cucullatus C. V.Morton, Contr. U.S. uador-Pastaza: Puyo, near the Rio Pastaza, Natl. Herb. 29: 36. 1944. TYPE: Guatema­ 850 m, 12 May 1935, H. Schultze-RhonhoJ la-Quiche: Zona Reyna, 750-900 m, A.F. 1850 (holotype: B, destroyed). TYPE: Ec­ Skutch 1812 (holotype: US!; isotypes: A, uador-Napo: Archidona, forest along the F!). =Glossoloma cucullatum (C.V.Morton) Baeza-Loreto road, 3-5 km E of the Tena­ J.L.Clark, comb. nov. Baeza junction, 27 Feb 2003, J.L. Clark & Alloplectus cucullatus C.VMorton var. substri­ N. Harris 7284 (neotype, here designated: gosus C.VMorton, Contr. U.S. Natl. Herb. QCNE!; isoneotypes: AAU! COL! E! GB! 29: 37. 1944. TYPE: Guatemala-Alta Ver­ MO! NY! SELl US!). =Drymonia crena­ tiloba (Mansf.) Wiehler, Selbyana 5(3,4): apaz: Pansamalli, 1150 m, Jun 1885, H. von 382. 1981. Tiierckheim 733 (holotype: US!; isotypes: Alloplectus cristatus (Griseb.) Fawc., Provo List. GH! NY! PH! US!). =Glossoloma cucul­ PI. Jam.: 28. 1893. Pterygoloma cristatum latum (C.V.Morton) J.L.Clark. Griseb., FI. Britt. W. I.: 464: 1862. Trichan­ Alloplectus darienensis Seem., Bot. Voy. Herald: tha cristata (Griseb.) Wi ehler, Selbyana 187. 1854. Episcia darienensis (Seem.) 1(1): 34. 1975. Ortholoma cristatum (Gri­ Leeuwenb., Acta. Bot. Need. 8: 52. 1959. seb.) Wi ehler, Phytologia 27: 321. 1973. =Paradrymonia darienensis (Seem.) =Columnea grisebachiana Kuntze, Revis. Wiehler, Selbyana 5(1): 50. 1978. Gen. PI. 2: 472. 1891. Alloplectus deltoideus C.VMorton, Fieldiana, Alloplectus cristatus (L.) Mart., Nov. Gen. Sp. Bot. 28: 521. 1953. =Corytoplectus deltoi­ PI. 3: 57. 1829 [text]. 3: 189. 1832 [index]. deus (C.V.Morton) Wiehler, Phytologia Besleria cristata L., Sp. PI.: 619. 1753. 27: 313. 1973. Prionoplectus cristatus (L.) Oerst., Centra­ Alloplectus dichrous (Spreng.) DC. Pro dr. 7: lamer. Gesner.: 78, pI. 9, 1858 [preprint); 546. 1839. Besleria dichrus Spreng. Syst. 152, pI. 9, 1861 [reprint). Columnea cristata Veg., ed. 16, 2: 840. 1825. Columnea di­ (L.) Kuntze, Revis. Gen. PI. 2: 472. 189l. chroa (Spreng.) Kuntze, Revis. Gen. PI. 2: Alloplectus cristatus (L.) Mart. var. typicus 472. 1891. Crantzia dichrus (Spreng.) Fournet, FI. Ill. Phan. Guad. Mart.: 1326. Fritsch in Engler & Prantl, Nat. Pflanzen­ 1978. =Crantzia cristata (L.) Scop., Intr. fam. 4(3b): 168. 1894. Nematanthus dichrus Hist. Nat.: 173. 1777. (Spreng.) Wiehler, Phytologia 27: 326. Alloplectus cristatus (L.) Mart. var. typicus 1973. =Nematanthus lanceolatus (Poir.) Fournet, Fl. Ill. Phan. Guad. Mart.: 1326. Chautems, Diss. Bot. 112: 189. 1988. 1978. =Crantzia cristata (L.) Scop., Intr. Alloplectus dichrous sensu Hook., Bot. Mag. 72: Hist. Nat.: 173. 1777. pI. 4216. 1846. =Nematanthus hirtellus CLARK: ALLOPLECTUS MONOGRAPH 201

(Schott) Wiehler, Baileya 18: 137. 1971 Alloplectus dimidiatus Benth., PI. Hartw.: 235. [1972]. 1846. Dalbergaria dimidiata (Benth.) Alloplectus dielsii (Mansf.) Wiehler, Selbyana Wiehler, Phytologia 27: 317. 1973. =Col­ 5(3,4): 381. Ortholoma dielsii (Mansf.) umnea dimidiata (Benth.) Kuntze, Revis. Wiehler, Phytologia 27: 321. 1973. Trichan­ Gen. PI. 2: 472. 1891. tha dielsii (Mansf.) Wiehler, Selbyana 1(1): Alloplectus dimorphobracteatus Hoehne, Sel­ 35.1975. TYPE: Ecuador-Chimborazo: Re­ lowia 9: 46. 1958. TYPE: Brazil-Amazon­ gion interandina. Tipococha, margen del as: Benjamin Constant, 18 Oct 1945, Murra bosque de montafia superior, 3200 m, 19 Pires & Black 922 (holotype: IAN, not lo­ Aug 1933, Diels 657 (holotype: B, de­ cated). It was not possible to verify the ex­ stroyed). TYPE: Ecuador-Carchi: cloud istence of the type specimen for this spe­ forest towards Maldonado, on road from cies, and it was not located by a previous Tulcan, 1800 m, 21 Feb 1978, C. Luer, J. visit to the IAN herbarium (A. Chautems Luer & A. Hirtz 2663 (neotype: SELl des­ pers. comm.). Its placement in Alloplectus ignated by Wiehler, Selbyana 5(3,4): 381- is dubious, because the species description 382. 1981). There is strong support for the is based on a single collection that lacked inclusion of Columnea dielsii in the genus flowers and was suspected as being new by Columnea from molecular and morpholog­ Hoehne, as it was unlike anything known ical data presented by Clark & Zimmer from the region where it was collected. Fur­ (2003) and Clark et ai. (in press). Its place­ thermore, it is described as a "cipo pe­ ment in Columnea is also congruent with queno" (small vine) with two types of results obtained by Smith and Sytsma bracts. There are no known small vines with (1994b) based on cpDNA restriction site two types of bracts in the current concept variation, where Columnea dielsii (voucher of Alloplectus, Crantzia, and Glossoloma. specimen determined as "Alloplectus peru­ Furthermore, the vague description of A. di­ vianus (Zahlbr.) L.P.Kvist & LE.Skog") morphobracteatus does not allow it to be was placed as a basal member of the genus placed in a known taxon. Columnea. This species was transferred Alloplectus discolor Hook.f., Bot. Mag. 118, in­ from Columnea to Alloplectus by Wiehler dex. 1892, nom. nud. This name appears in (1981). Columnea dielsii also was recog­ the index incorrectly as plate 7271 Allo­ nized as a heterotypic synonym of Alloplec­ plectus discolor. Plate 7271 corre'6ponds to tus peruvianus by Kvist & Skog (1993). Alloplectus lynchei Hook.f. =Nautilocalyx Wiehler (1981) recognized this species as a lynchii (Hook.f.) Sprague, Bull. '.\1isc. In­ member of Alloplectus because of the pres­ form. 1912: 89. 1912. ence of a capsular fruit. The inclusion of Alloplectus dodsonii Wiehler, Selbyana 2(1): 67, this taxon in Columnea is an anomaly, be­ pI. 19A. 1977. TYPE: Ecuador-Los Rios, cause it is one of only two known species Rio Palenque Science Center, km 56, be­ in the genus with a capsular fruit instead of tween Quevedo and Santo Domingo, 200 m, a berry. The position of Columnea dielsii, 19 Jul 1971, H. Wiehler & c.H. Dodson however, is not surprising, because it shares 7105 (holotype: SELl). =Drymonia dod­ many features with other members of Col­ sonii (Wiehler) J.L.Clark, comb. nov. umnea such as a 5-lobed nectary, non-re­ Alloplectus domingensis Urb. Symb. Antill. 2: supinate flowers, a scandent habit, and flow­ 357. 1901. Ortholoma domingense (Urb.) ers that are weakly bilobed (e.g., members Wiehler, Phytologia 27: 321. 1972. Trichan­ of Columnea sect. Pentadenia). It is not tha domingensis (Urb.) Wiehler, Selbyana clear why Wiehler (1973) initially treated 1(1): 34. 1975. =Columnea domingensis this species as a member of Ortholoma and (Urb.) B.D.Morley, Proc. Roy. Irish Acad. then transferred it to Trichantha (Wiehler 74B: 412, 424. 1974. 1975) instead of Pentadenia, where a 5- Alloplectus domingensis Urb. var. microphyllus lobed nectary is a defining feature. Never­ C.V.Morton, Contr. U.S. Nati. Herb. 28(1): theless, the presence of a 5-lobed nectary is 19. 1944. =Columnea domingensis (Urb.) not known to exist in any species of Allo­ B.D.Morley, Proc. Roy. Irish Acad. 74B: plectus sensu stricto. The only other genus 412, 424. 1974. of Episcieae besides Columnea with a 5- Alloplectus doratostylus Leeuwenb., Acta Bot. lobed nectary gland is Corytoplectus Oerst., Need. 8: 44. 1959. =Drymonia doratostyla and most genera in the tribe Episcieae have (Leeuwenb.) Wiehler, Phytologia 27: 325. a bilobed gland or two large dorsal connate 1973. glands. =Columnea dielsii Mansf., Bib­ Alloplectus eriocalyx Galeotti in J.D. Hooker, lioth. Bot. 116: 145. 1937. Bot. Mag. 74: pI. 4371. 1848, nom. nud. 202 SELBYANA Volume 25(2) 2005

This name was attributed to the specimen (holotype: US!; isotype: F!). =Glossoloma sent to Hooker by Galeotti in 1846, which pycnosuzygium (Donn.Sm.) J.L.Clark. was grown at Kew and used for making the Alloplectus grandicalyx J.L.Clark & L.E.Skog, plate of Alloplectus concolor Hook.f. Thus, Novon 12: 173-178. 2002 (Clark & Skog this name was mentioned in the text, but not 2002). TYPE: Ecuador-Napo: Canton Ar­ used as the specific epithet for the plate of chidona, Reserva Ecologica Antisana, Sec­ A. concolor Hook.f. =Nematanthus lan­ tor Guacamayos, Sendero Jumandy-La Vir­ ceolatus (Poir.) Chautems, Diss. Bot. 112: gen (km 30), path that follows potential oil 189. 1988. pipeline, 00038'S, 77°49'W, 1700-1800 m, Alloplectus erythroloma Leeuwenb., Acta Bot. 1 May 1998, l.L. Clark. E. Narvaez. & T. Neerl. 8: 45. 1959. =Drymonia erythrolo­ Pauchi 5449 (holotype: QCNE!; isotypes: ma (Leewenb.) Wiehler, Selbyana 2(1): AAU! COL! E! MO! QCA! SRP! US!). 74, pI. 22A. 1977. =Glossoloma grandicalyx (J.L.Clark & Alloplectus forgetii Sprague. Bull. Misc. Inform. L.E.Skog) J.L.Clark, comb. nov. Kew 1910: 383. 1910. =Nautilocalyx for­ Alloplectus grandifoUus Britton ex Rusby, Bull. getii (Sprague) Sprague, Bull. Misc. In­ Torrey Bot. Club 27: 30. 1900. =Cory to­ form. Kew 1912: 89. 1912. plectus riceanus (Rusby) Wiehler, Phyto­ Alloplectus forseithii (Oerst.) Hanst., Linnaea logia 27: 313. 1973. 34: 375. 1865. Prionoplectus forseithii Alloplectus grisebachianus (Kuntze) Urb. Symb. Oerst., Centralamer. Gesner.: 45, pI. 9 (figs. Antill. 2: 357. 1901. =Columnea griseba­ 30-35), 1858 [preprint]; 119, pI. 9 (figs. chiana Kuntze, Revis. Gen. PI. 2: 472. 30-35), 1861 [reprint]. =Crantzia cristata 1891. (L.) Scop., Intr. Hist. Nat.: 173. 1777. Alloplectus grisebachianus (Kuntze) Urb. Var' Alloplectus fraternus C.V.Morton, Proc. BioI. ochrotrichus Urb. Symb. Antill. 5: 497. Soc. Wash. 69: 193. 1956. TYPE: Panama­ 1908. =Columnea grisebachiana Kuntze, Chiriqui: flat areas on the slopes of Volcin Revis. Gen. PI. 2: 472. 1891. de Chiriqui Viejo and along Rio Chiriqui Alloplectus guatemalensis C. V, Morton, Contr. Viejo, 1200 m, 20 Apr 1935, Allen 1000 U.S. Natl. Herb. 29: 37. 1944. =Drymonia (holotype: MO! fragment: US!; isotypes: guatemalensis (C.V.Morton) D.N.Gibson, BM! BR! F! G! U!). =Glossoloma tetra­ Phytologia 23: 336. (Gibson 1972). Alloplectus guianensis (C.V.Morton) Stearn, gonum Hanst., Linnaea 26: 208-209. 1854. Bull. Brit. Mus. (Nat. Hist.) Bot. 4: 189. Alloplectus glaber (Hook.f.) Decne., Rev. Hort. 1969. Dalbergaria guianensis (C.V,Morton) 21: 243. 1849. Hypocyrta glabra Hook.f., Wieh1er, Phytologia 27: 317. 1973. =Col­ Bot. Mag. 74: pI. 4346. 1848. =Nematan­ umnea guianensis C.V.Morton, Bull. Tor­ thus strigillosus (Mart.) H.E.Moore, Bail­ rey Bot. Club 75: 564. 1948. eya 19: 38. 1973, fide Chautems (1988). Alloplectus herthae Mansf., Notizbl. Bot. Gart. Alloplectus glaber DC. Prodr. 7: 546. 1839. LEC­ Berlin-Dahlem 14: 38. 1938. TYPE: Ecua­ TOTYPE: Sesse and Mocifio plate, number dor-"San Carlos" [=San Carlos de los 6331.633 in the Tomer Collection of Sesse Colorados], 120 m, 8 Sept 1935, H. Schul­ and Mocifio Biological Illustrations at the tze-Rhonhof 1903 (holotype: B, de­ Hunt Institute for Botanical Documentation. stroyed).-Pichincha: along old road be­ McVaugh (2000) lectotypified this name tween Santo Domingo-Quito (via Chiribo­ with the following notes linking this plate ga), 1400-1600 m, 28 Oct 1995, P. Men­ to the de Candolle description, "cited in doza-T. et al. 618 (neotype, here synonymy by DC., I.c., as the basis for Al­ designated: QCNE!; isoneotypes: AAU! loplectus ? glaber. Besleria ["Besteria"] CAS! COL! E! F! GB! MO! NY! QCA! cristata [L.] sensu Sesse & Mocifio PI. Nov. SRP! US!). =Glossoloma herthae (Mansf.) Hisp. 103. 1889; ed. 2. 97. 1893." Locality J.L.Clark, comb. nov. notes from de Candolle include: Mountains Alloplectus hirsutus Sprague, Bull. Misc. In­ of Mazatlin, Guerrero." =Drymonia ser­ form. Kew 1911: 346.1911. =Nautilocalyx rulata (Jacquin) Mart., Nov. Gen. Sp. PI. hirsutus (Sprague) Sprague, Bull. Misc. 3: 59 [text]. 1829.3: 189 [index]. 1832. Inform. 1912: 89. 1912. Alloplectus glabrescens C.Y.Morton, Gesneri­ Alloplectus hirtellus (Schott) Preston ex Hoehne, aceae in J.A. Steyermark, Fieldiana, Bot. Sellowia 9: 45. 1958. =Nematanthus hir­ 28: 522. 1953. TYPE: Venezuela-Merida: tellus (Schott) Wiehler, Baileya 18: 137: north of Torondoy, along Quebrada Molino, 1971 [1972]. in woods above Las Cuadras, 1820-2255 Alloplectus hirtellus (Schott) Preston ex Hoehne m, 27 Mar 1944, l.A. Steyermark 55818 forma crassifolius Hoehne, Sellowia 9: 45. CLARK: ALLOPLECTUS MONOGRAPH 203

1958. =Nematanthus hirtellus (Schott) 186. 1854. Dalbergaria lanata (Seem.) Wiehler, Baileya 18: 137: 1971 [1972]. Wiehler. =Columnea lanata (Seem.) Kunt­ Alloplectus hirtellus (Schott) Preston ex Hoehne ze, Revis. Gen. Pl. 2: 472. 1891. forma umbrophilus Hoehne, Sellowia 9: 45. Alloplectus lindenii (Kuntze) Wiehler, Phytolo­ 1958. =Nematanthus hirtellus (Schott) gia 27: 327. 1973. Crantzia lindenii (Kun­ Wiehler, Baileya 18: 137: 1971 [1972]. tze) Fritsch in Engler & Prantl, Nat. Pflan­ Alloplectus hoppii Mansf., Repert. Spec. Nov. zenfam. 4(3b): 168. 1894. Columnea lin­ RegniVeg.41: 148. 1936. TYPE: Ecuador-­ denii Kuntze, Revis. Gen. PI. 2: 471. 1891. Napo: Archidona, Jan 1931, W. Hopp 1032 =Alloplectus weirii (Kuntze) Wiehler, (syntype: B, destroyed); Tena: hacienda dos Phytologia 27: 327. 1973. Rios, 8 Apr 1933, E. Heinrichs 319 (syn­ Alloplectus lynchei Hook.f., Bot. Mag. 118: pI. type: B, destroyed); Pastaza: near the Rio 7271. 1892. Episcia lynchei (Hook.f.) Pastaza, 12 May 1935, H. Schultze-Rhonhof Kunth, Repert. Spec. Nov. Regni Veg. Beih. 1838 (syntype: B, destroyed). TYPE: Ecua­ 43: 643. 1928. =Nautilocalyx lynchii dor--Napo: Cant6n Tena, Parroquia Talag, (Hook.f.) Sprague, Bull. Misc. Inform. Comunidad Cando, N or Rio Jatunyacu, 540 1912: 89. 1912. m, 1°4'45"S, 77°56'29"W, 21 Dec 2000, J.L. Alloplectus macranthus Donn.Sm., Bot. Gaz. 31: Clark, E. Narvaez & V. Grefa 5669 (neo­ 117. 1901. =Drymonia macrantha (Donn. type, here designated: QCNE!; isoneotypes: Sm.) D.N.Gibson, Phytologia 23: 336. AAU! COL! NY! QCA! US!). =Drymonia 1972. hoppii (Mansf.) Wiehler, Phytologia 27: Alloplectus macrophyllus (Oerst.) Hemsi. in 325. 1973. Godman & Salvin, BioI. Cent.-Amer., Bot. Alloplectus ichthyodenna Hanst., Linnaea 34: 2: 484. 1882. Caloplectus macrophyllus 372. 1865. Columnea ichthyoderma Oerst. Centralamer. Gesner.: 45, pI. 9 (figs. (Hanst.) Kuntze, Revis. Gen. PI. 2: 472. 44-46), 1858 [preprint]; 119, pI. 9 (figs. 1891. Crantzia ichthyoderma (Hanst.) 44-46), 1861 [reprint]. Columnea macro­ Fritsch in Engler & Prantl, Nat. Pflanzen­ phylla (Oerst.) Kuntze, Revis. Gen. PI. 2: fam.4(3b): 168. 1894. TYPE: Costa Rica: H. 472. 1891. =Drymonia macrophylla (Oerst.) Wendland 833 (holotype: GOET!). =Glos­ H.E.Moore, Baileya 3: 112. 1955. soloma ichthyoderma (Hanst.) J.L.C1ark, Alloplectus martinianus J.RSmith, Novon 9: comb. nov. 419. 1999. TYPE: Ecuador--Napo: road Alloplectus ichthyodenna Hanst. var. hirsutulus from Baeza to Tena, 5 km S of Baeza, 26 C.V.Morton, PubI. Field Mus. Nat. Hist., Jun 1983, J. Bohlin & M. Bohlin 698 (ho­ Bot. Ser. 18: 1145. 1938 (Morton 1938). lotype: US!; isotype: GB). =Glossoloma TYPE: Costa Rica--Cartago: Atirro, 600 m, martinianum (J.F.Smith) J.L.Clark, Apr 1896, J. Donnell Smith 6715 (holotype: comb. nov. US!; isotypes: BM! GH! K! US!). =Glos­ Alloplectus medusaeus L.E.Skog, Ann. Missouri soloma tetragonum Hanst., Linnaea 26: Bot. Gard. 65: 797. 1979. TYPE: Panama-­ 208-209. 1854. Chiriqui: on slope of La Popa above Bo­ Alloplectus ichthyoderma var. pallidus C. V. quete, 1600 m, 5 Aug 1972, w.G. D'Arcy Morton, PubI. Field Mus. Nat. Hist., Bot. & J.J. D'Arcy 6398 (holotype: US!; iso­ Ser. 18: 1145. 1938. TYPE: Costa Rica--San types: MO-2 sheets! SELl). =Glossoloma Jose: Las Nubes, 1500-1900 m, 20-22 Mar medusaeum (L.E.Skog) J.L.Clark, comb. 1924, P.e. Standley 38793 (holotype: US!). nov. =Glossoloma ichthyoderma (Hanst.) Alloplectus melittifolius Mart. ex Loudon, En­ J.L.Clark. cycI. PI. new ed.: 1402. 1855, nom. nud. pro Alloplectus ichthyodenna forma rub esc ens syn. =Nautilocalyx melittifolius (L.) C.V.Morton, Pubi. Field Mus. Nat. Hist., Wiehler, Phytologia 27: 307. 1973. Bot. Ser. 18: 1145. 1938. TYPE: Costa Alloplectus meridensis Klotzsch ex Hanst., Lin­ Rica--Cartago: Santa Clara, 1950 m, 20 Jul naea 34: 372. 1865. Columnea meridensis 1923, W.R. Maxon & A.D. Harvey 8195 (Klotzsch ex Hanst.) Kuntze, Revis. Gen. (holotype: US I). =Glossoloma ichthyoder­ PI. 2: 472. 1891. TYPE: Venezuela--Merida. ma (Hanst.) J.L.Clark. J. Moritz 1132 (holotype: B, destroyed; lec­ Alloplectus kunthii G.Don, Gen. Hist. 4: 655. totype: here designated, G!; isolectotypes: 1837 [1838]. Don created this superfluous BM, BR! L, PI). Recent authors (Smith & name instead of using Alloplectus hicolor Sytsma 1994a) have used this name on (Kunth) Klotzsch ex Hanst. =Glossoloma voucher specimens for molecular systematic bicolor (Kunth) J.L.Clark. studies. =Glossoloma ichthyoderma (Hanst.) Alloplectus lanatus Seem., Bot. Voy. Herald: J.L.Clark. 204 SELBYANA Volume 25(2) 2005

Alloplectus metamorphophyllus Donn.Sm., Bot. paramo and ridge top, by boundary line Gaz. 52: 52. 1911. TYPE: Costa Rica-San Lara-Trujillo, above Humocaro Bajo, 2600- Jose: La Palma, 1600 m, A. Tonduz 10884 2800 m, 6 Feb 1944, J.A. Steyermark 55355 (lectotype, designated by Wiehler, Selbyana (holotype: US!; isotypes: F! NY! VEN!). 5: 57. 1978: US!). =Paradrymonia meta­ =Glossoloma chrysanthum (Planch. & morphophylla (Donn.Sm.) Wiehler, Phy­ Linden) J.L.Clark. tologia 27: 327. 1973. Alloplectus pallidus Sprague, Bull. Misc. In­ Alloplectus microsepalus C.Y.Morton, Fieldiana, form. 1911: 346. 1911. =Nautilocalyx pal­ Bot. 28: 523. 1953. Columnea microsepala lidus (Sprague) Sprague, Bull. Misc. In­ (C.V.Morton) L.P.Kvist & L.E.Skog, Aller­ form. 1912: 89. 1912. tonia 6: 391. 1993. Pentadenia microsepala Alloplectus panamensis C.Y.Morton, Ann. (C.Y.Morton) Wiehler, Phytologia 27: 315. Missouri Bot. Gard. 29: 36. 1942. TYPE: 1973. TYPE: Venezuela-Monagas. Cerro Panama-Cocle: Hills north of EI Valle de de la Cueva de Dona Anita, S of Caripe, Anton, 1000 m, 14 Jul 1940, P.H. Allen 1100-1200 m, 7 Apr 1945, J.A. Steyermark 2189 (holotype: US!; isotypes: MO! S!). 61905 (holotype: F; isotypes: NY, US!). =Glossoloma panamense (C.V.Morton) =Columnea spathulata Mansf., NotizbI. J.L.Clark, comb. nov. Bot. Gart. Berlin. 14: 37. 1838, fide Smith Alloplectus parviflorus (Hanst.) Hemsi. in God­ (1994). man & Salvin, BioI. Cent.-Amer., Bot. 2: Alloplectus multiflorus (Oerst. ex Hanst.) Hanst., 484. 1882. Polythysania parviflora Hanst., Linnaea 34: 361. 1865. Calanthus multiflo­ Linnaea 26: 209, pI. 55. 1854. =Drymonia rus Oerst. ex Hanst., Linnaea 26: 208-209. hansteiniana Wiehler, Phytologia 27: 325. 1854. Columnea multiflora (Oerst. ex 1973. Hanst.) Kuntze, Revis. Gen. Pl. 2: 472. Alloplectus parviflorus Hanst., Linnaea 34: 377, 1891. Crantzia multiflora (Oerst. ex Hanst.) 1865. Crantzia parviflora (Hanst.) Fritsch Fritsch, Fritsch in Engler & Prantl, Nat. I in Engler & Prantl, Nat. Pflanzenfam. 4(3b): Pflanzenfam. 4(3b): 168. 1894. A detailed 168. 1894. Hanstein published this name description of this species can be found in and referenced the Mart. 1829 species de­ Oersted (Centralamer. Gesner.: 47, pI. 3, scription of Alloplectus sparsiflorus Mart. 1858 [preprint]; 121, pI. 3, 1861 [reprint]). The basionym for this name that appears in Oersted's combination and a section of his Hanstein (1865) is therefore an error for AI­ illustration were published by Hanstein. Al­ loplectus sparsiflorus Mart. =Nematanthus though Hanstein attributed this name to hirtellus (Schott) Wiehler, Baileya 18: Oersted, it had not been priorly published 137. 1971 [1972]. and therefore is treated as validated by Han­ Alloplectus patrisii DC. Prodr. 7: 545. 1839. Ma­ stein. =Drymonia multiflora (Oerst. ex crochlamys patrisii (DC.) Decne., Rev. Hanst.) Wiehler, Phytologia 27: 325. 1973. Hort. 21: 243. 1849. Columnea patrisii Alloplectus nummularia (Hanst.) Wiehler, Bail­ (DC.) Kuntze, Revis. Gen. PI.: 2. 1891. eya 18: 136. 1971 [1972]. Hypocyrta num­ Crantzia patrisii (DC.) Fritsch in Engler & mularia Hanst., Linnaea 34: 381. 1865. Col­ Prantl, Nat. Pflanzenfam. 4(3b): 168. 1894. umnea nummularia (Hanst.) Kuntze. =Neo­ =Drymonia coccinea (Aubl.) Wiehler, mortonia nummularia (Hanst.) Wiehler, Phytologia 27: 324. 1973. Selbyana 5(1): 63. 1978. Alloplectus patrisii Miquel ex Decne. in Rev. Alloplectus oblongicalyx J.L.Clark & L.E.Skog, Hort. 21: 243. 1849, nom. nud. pro syn. Novon 15: 74. 2005. TYPE: Ecuador-Pi­ Macrochlamys miquelii Decne., Rev. Hort. chincha: Canton Quito, trail betw. Uoa & 21: 243. 1849, nom. nud. Decaisne differ­ Mindo, 30 km W of Uoa, OOol1'S, entiated de Candolle's homonym of Mi­ 78°41'W, 1800 m, 12 Feb 1997, J.L. Clark quel's name by, "Macrochlamys miquelii & S.c. Clark 3903 (Holotype: QCNE!; iso­ Decne. =Alloplectus patrisii Miquel, non types: AAU! MO! NY! US!). =Glossoloma DC." Decaisne's generic description of Ma­ oblongicalyx (J.L.Clark & L.E.Skog) crochlamys was too vague for assigning this J.L.Clark, comb. nov. name with certainty to genus, and there are Alloplectus oinochrophyllus Donn.Sm., Bot. no known annotated herbarium collections Gaz. 54: 239. 1912. =Drymonia oinoch­ with either of these names. In the absence rophylla (Donn.Sm.) D.N.Gibson, Phyto­ of information, it is not possible to assign logia 23: 336. 1972. this name with certainty. Alloplectus ornatus C.V.Morton, Fieldiana, Bot. Alloplectus peltatus Oliver, Bot. Mag. 103: pI. 28: 523. 1953. TYPE: Venezuela-Trujillo: 6333. 1877. Columnea peltata (Oliver) La Quebrada Cortijo, in dense woods below Kuntze, Revis. Gen. PI. 2: 472. 1891. CLARK: ALLOPLECTUS MONOGRAPH 205

=Drymonia peltata (Oliver) H.E.Moore, m, 2 May 1982, B. 011gaard et al. 37673 Baileya 3: 112. 1955. (holotype: AAUl). =Glossoloma purpu­ Allopleetus penduliflorus M.Freiberg, Phyton reum (L.P.Kvist & L.E.Skog) J.L.Clark, 37: 133. 1997. TYPE: Ecuador-Imbabura: comb. nov. Cant6n Sigchos, Los Cedros Biological Re­ Allopleetus pyenosuzygius (Donn. Sm.) C.Y. serve, 1200-1500 m, 27 Jun 1996, M. Frei­ Morton, Fieldiana, Bot. 28: 522. 1953. Bes­ berg 96210 (holotype: QCA; isotypes: Ie ria pyenosuzygia Donn.Sm., Bot. Gaz. 52: ULM-3 sheets!). =Glossoloma penduliflo­ 53. 1911. TYPE: Costa Rica-San Jose, for­ rum (M.Freiberg) J.L.Clark, comb. nov. ests at La Palma, 1459 m, 8 Sep 1898, A. Allopleetus pendulus Poepp. in Poeppig & End­ Tonduz 12545 (lectotype, here designated, licher, Nov. Gen. Sp. PI. 3: 6, pI. 205. 1840. US!; isolectotypes: CRt F! K! SEL! US!). Crantzia pendula (Poepp.) Fritsch in Engler =Glossoloma pycnosuzygium (Donn.Sm.) & Prantl, Pflanzenfam. 4(3b): 168. 1894. J.L.Clark, comb. nov. =Drymonia pendula (Poepp.) Wiehler, Allopleetus repens Hook., Bot. Mag. 72: pI. Phytologia 27: 325. 1973. 4250. 1846. Pterygoloma repens (Hook.) Allopleetus peruvianus (Zahlbr.) L.P.Kvist & Hanst., Linnaea 26: 211. 1854. Ortholoma L.E.Skog, Allertonia 6(5): 295. 1993. Or­ repens (Hook.) Wiehler, Phytologia 27: 323. tholoma peruvianum (Zahlbr.) Wiehler. Phy­ 1973. =Columnea repens (Hook.) Hanst., tologia 27: 322. 1973. Triehantha peruvi­ Linnaea 34: 396. 1865. ana (Zahlbr.) Wiehler, Selbyana 1(1): 35. Allopleetus ruaeophilus Donn.Sm., Bot. Gaz. 1975. This name has been incorrectly attri­ 54: 238. 1912. TYPE: Costa Rica-Heredia: buted in the literature to plants from Ecua­ Volcan de Barba, 2500-2700 m, Feb 1890, dor (Kvist & Skog 1993, Smith & Sytsma A. Tonduz 1997 (lectotype designated by 1994b, Skog & Kvist 1997) as well as on C.Y. Morton (1938): US!; isolectotypes: herbarium specimens. The correct name for BR! U! WI). =Glossoloma ichthyoderma the material from Ecuador is Columnea (Hanst.) J.L.Clark. dielsii Mansf. (see above under A. dielsii). Allopleetus rubidus C. V.N1orton, Ann. Missouri Based on fieldwork and studies of type ma­ Bot. Gard. 24: 204. 1937. Pentadenia rub­ terial of these two taxa, it is apparent that ida (C.Y.Morton) Wiehler, Selbyana 2(1): they represent different species. The type 122. 1977. =Columnea rubida (C.V. collection from Peru (treated here as Col­ Morton) C.V.Morton, Baileya 7: 58. 1959. umnea peruviana Zahlbr.) represents a spe­ Allopleetus sanguineus (Pers.) G.Don, Gen. cies with a baccate fruit, and the type col­ Hist. 4: 655. 1837 [1838]. Besleria sangui­ lection from Ecuador (treated here as Col­ nea Pers., Syn PI. 2: 165. 1807. Collandra umnea dielsii Mansf.) represents a species sanguinea (Pers.) Griseb., Mem. Amer. with a capsular fruit. TYPE: Peru-Tambillo, Acad. Arts 8: 526. 1861 [1862]. Dalbergar­ C. felski 72 (holotype: WI). =Columnea ia sanguinea (Pers.) Steudel, Nomencl. Bot. peruviana Zahlbr., Ann. K. K. Naturhist. Ed. 2, 1: 479. 1849. =Columnea sanguinea Hofmus. 7: 8. 1892. (Pers.) Hanst., Linnaea 34: 384. 1865. Allopleetus pinelianus Lem., Fl. Serres Jard. Eur. Allopleetus savanna rum C.V.Morton, Bull. Tor­ 2(8): pI. 5. 1846. =Nematanthus lanceo­ rey Bot. Club 75: 563. 1948. TYPE: Guya­ latus (Poir.) Chautems, Diss. Bot. 112: na-Kaieteur Plateau: savanna, wallaba for­ 189. 1988 (Chautems 1988). est [Eperua spp.], 1 May 1944, B. Maguire Allopleetus pinelianus Lem. var. bieolor Lem., & D.E. Fanshawe 23127 (holotype: NY!; FI. Serres Jard. Eur. 2(8): sub pI. 5. 1846. isotypes: A (photo: US!), BR! F! G! K! MO =Nematanthus lanceolatus (Poir.) Chau­ (photo: US!), P, Ul UC! US! VEN! WI). tems, Diss. Bot. 112: 189. 1988. =Nematanthus savannarum (C.V.Morton) Allopleetus pubeseens (Griseb.) Fawc., Provo J.L.Clark, comb. nov. List PI. Jam.: 28. 1893. Pterygoloma pu­ Allopleetus seabridus Lem., Jard. Fleur. 1: Misc. beseens Griseb., FI. Brit. W. I.: 464: 1862. 51. Hypoeyrta seabrida Lem., Fl. Serres Ortholoma pubeseens (Griseb.) Wiehler, Jard. Eur. 3: 238, pI. 6. 1847. 1851. Col­ Phytologia 27: 323. 1973. Triehantha pu­ umnea seabrida (Lem.) Kuntze, Rev. Gen. beseens (Griseb.) Wiehler, Selbyana 1(1): FI. 2: 472. 1891. =Nematanthus strigillo­ 35. 1975. =Columnea pubescens (Griseb.) sus (Mart.) H.E.Moore, Baileya 19: 38. Kuntze, Revis. Gen. PI. 2: 472. 1891. 1973, fide Chautems (1988). Allopleetus purpureus L.P.Kvist & L.E.Skog, Allopleetus sehlimii Planch. & Linden, FI. Serres Brittonia, 44: 475. 1992. TYPE: Ecuador­ Jard. Eur. 8: 211, pI. 827. 1853. Columnea Pichincha: Cant6n Quito, road EI Paraiso­ sehlimii (Planch. & Linden) Kuntze, Revis. Saguangal, km 11, 0012'N, 78°46'W, 1200 Gen. PI. 2: 472. 1891. =Corytoplectus 206 SELBYANA Volume 25(2) 2005

schlimii (Planch. & Linden) Wiehler, 75: pI. 4452. 1849, nom. nud. pro syn. Phytologia 27: 313. 1973. Hooker cited this name as a plant in culti­ Alloplectus schottii G.Don, Gen. Hist. 4: 655. vation in the protologue of Alloplectus cap­ 1837 [1838]. Don published Alloplectus itatus, and therefore it is presumed that this schottii as a new name and cited Besleria name appears in Hooker's publication as a dichrus Spreng. and Besleria bicolor Schott synonym of Alloplectus capitatus Hook. as belonging to this species. =:;Nematanthus =Corytoplectus capitatus (Hook.) Wieh­ lanceolatus (Poir.) Chautems, Diss. Bot. ler, Phytologia 27: 313. 1973. 112: 189. 1988. Alloplectus speciosus Poepp. in Poeppig & End­ Alloplectus schultzei Mansf., Repert. Spec. Nov. licher, Nov. Gen. Sp. Pl. 3: 6. 1840. Crant­ Regni Veg. 36: 124. 1934. TYPE: Col om­ zia speciosa (Poepp.) Fritsch in Engler & bia-Cundinamarca: mountain jungle near Prantl, Pfianzenfam. 4(3b): 168. 1894. Aguadita [5 km NE of Fusagasuga], 12 Apr =Corytoplectus speciosus (Poepp.) Wieh­ 1925. A. Schultze 212 (holotype: B, de­ ler, Phytologia 27: 313. 1973. stroyed). TYPE: Colombia-Cundinamarca: Alloplectus spectabilis Wiehler ex L.E.Skog & mountains W of SaIto de Tequendama, La­ Steyerm., Opera Bot. 92: 228. 1987. TYPE: guna de Catarnica, S of Santandercito, 2000 Venezuela-Sucre: Peninsula de Paria, Cer­ m, 31 Oct 1959, L. Uribe U. 3414 (neotype, ro Patao, N of Puerto de Hierro, NE of Gui­ here designated: US i). =Glossoloma ria, cloud forest on fiat mountain summit, schultzei (Mansf.) J.L.Clark, comb. nov. 1020 m, 19 Jul 1962, J.A. Steyermark & G. Alloplectus semicordatus Poepp. in Poeppig & Agostini 91094 (holotype: US!; isotypes: F! Endlicher, Nov. Gen. Sp. PI. 3: 5. 1840. NY! VEN!). =Crantzia spectabilis (Wieh­ Columnea semicordata (Poepp.) Kuntze, ler ex L.E.Skog & Steyerm.) J.L.Clark, Revis. Gen. PI. 2: 472. 1891. Crantzia sem­ comb. nov. icordata (Poepp.) Fritsch in Engler & Alloplectus splendens G.Don in Loudon, EncycI. Prantl, Pflanzenfam. 4(3b): 168. 1894. PI. new ed.: 1402. 1855, nom. nud. There =Drymonia semicordata (Poepp.) Wieh­ are no known annotated specimens with this ler, Phytologia 27: 326. 1973. name, and it does not appear in Don (1838). Alloplectus serpens J.L.Clark & L.E.Skog, No­ It is cited in Loudon as belonging to the von 15: 76. 2005. TYPE: Ecuador-Morona­ same genus as Hypocyrta discolor Lindley Santiago: Canton Limon-Indanza, main (= Nematanthus hirtellus (Schott) Wiehler). road betw. Gualaceo & Plan de Milagro, In the absence of a detailed description and "Tinaji1las, " cow pasture & patches of lack of locality information, it is not possi­ montane forest, 03°00' 19"N, 78°36'36"W, ble to assign this name to a species. 2800 m, 10 Jan. 2001, J.L. Clark, F. San­ Alloplectus ~prucei (Kuntze) Wiehler, Phytolo­ chez, & L. Jost 5904 (hoJotype: QCNE!; isotypes: AUU! HA! K! MO! NY! QCA! gia 27: 327. 1973. Columnea sprucei Kun­ US i). =Glossoloma serpens (J.L.Clark & tze, Revis. Gen. PI. 2: 471. 1891. Crantzia L.E.Skog) J.L.Clark, comb. nov. sprucei (Kuntze) Fritsch in Engler & Prantl, Alloplectus simulatus C.Y.Morton, Ann. Missou­ Nat. Pfianzenfam. 4(3b): 168. 1894. TYPE: ri Bot. Gard. 29: 37. 1942. TYPE: Panama­ Ecuador-Bolivar: El Limon, at the base of Chiriqui: vicinity of Bajo Mona and Que­ the mountain Chimborazo, ca. 900 m, Jun brada Chiquero, 1500 m, 18 Jul 1940, R.E. 1860, R. Spruce 6216 (holotype: K!; iso­ Woodson & R. W. Schery 537 (holotype: types: BM! W [photo: US!]). =Glossoloma US!; isotypes: GH [photo: US!], MO!). sprucei (Kuntze) J.L.Clark, comb. nov. =Glossoloma tetragonum Hanst., Linnaea Alloplectus stenophyllus Donn.Sm., Bot. Gaz. 26: 208-209. 1854. 25: 153. 1898. =Drymonia stenophylla Alloplectus sparsifiorus Mart., Nov. Gen. Sp. PI. (Donn. Sm.) H.E.Moore, Baileya 3: 112. 3: 55, pI. 223, fig. 1. 1829. Columnea spar­ 1955. s(flora (Mart.) Kuntze, Revis. Gen. PI. 2: Alloplectus strigosus (Oerst.) Hanst., Linnaea 472. 1891. Crantzia sparsifiora (Mart.) Re­ 34: 374. 1865. Saccoplectus strigosus chinger, Oersterr. Bot. Zeit. 49: 18l. 1899. Oerst., Centralamer. Gesner.: 44, 1858 [pre­ =Nematanthus hirtellus (Schott) Wiehier, print]; 118, 1861 [reprint]. Crantzia strigo­ Baileya 18: 137. 1971 [1972]. sa (Oerst.) Fritsch in Engler & Prantl, Nat. Alloplectus solitarius Rusby, Mem. Torrey Bot. Pfianzenfam. 4(3b): 168. 1894. =Drymonia Club 4: 238. 1895. =Drymonia solitaria strigosa (Oerst.) Wiehler, Baileya 18: 137. (Rusby) Wiehler, Phytologia 27(5): 326. 1971 [1972]. 1973. Alloplectus sylva rum S.Moore, Trans. Linn. Soc. Alloplectus speciosus Hort. ex Hook., Bot. Mag. London, Bot. Ser. 2 4: 411. 1895. =Dry- CLARK: ALLOPLECTUS MONOGRAPH 207

monia semicordata (poepp.) Wiehler, Phy­ Fritsch in Engler & Prantl, Nat. Pflanzen­ ! tologia 27: 326. 1973. fam 4(3b): 168. 1894. TYPE: PI. 11 in Kar­ AUoplectus tenuis Benth., PI. Hartw.: 235. 1846. sten, Ausw. Gew. Venez. 34-36, pI. 11. Columnea benthamiana Kuntze, Revis. 1848. No authentic Karsten collections of Gen. PI. 2: 471. 1891, non Columnea tenuis this species have been located or cited in Klotzsch ex Oerst. TYPE: Ecuador: K. Har­ the publication. Karsten cites the type lo­ tweg s.n. (holotype: K!). =Drymonia tenuis cality as near Caracas (Venezuela). Most (Benth.) J.L.Clark, comb. nov. collections of this species are from Colonia A(loplectus tetragonoides Mansf., Biblioth. Bot. Tovar, in the Estado Aragua of Venezuela. I 116: 144-145. 1937. TYPE: Ecuador-TIm­ =Crantzia tigrina (Karst.) J.L.Clark, gurahua: area of Rio Pastaza and Rio Topo, comb. nov. 1200 m, 12 Sept 1933, L. Diels 1002 (B, no Alloplectus tomentosus Glaz., Bull. Soc. Bot. longer extant). TYPE: Ecuador-Tungura- France, Mem. 3: 515. 1909 [1911], nom. I hua: Cant6n Banos, Parroquia Rio Negro, nud. = Vanhouttea lanata Fritsch, Bot. I near Rio Topo, 1400 m, 17 Jan 2001, J.L. Jahrb. Syst. 29 (Beibl. 65): 16. 1900, fide Clark & V. Duran 6000 (neotype, designat­ Wiehler (1973). ed by J.L. Clark & L.E. Skog (2002): Alloplectus tucurriquensis Donn.Sm., Bot. Gaz. QCNE!; isoneotypes; AAU! COL! F! K! 54: 238. 1912. TYPE: Costa Rica-Cartago: MO! NY! QCA! S! US!). =Glossoloma te- Las Vueltas, Tucurrique, 635-700 m,

I tragonoides (Mansf.) J.L.Clark, comb. 9°51'N, 83°43'W, Mar 1899, A. Tonduz ! nov. 13042 (holotype: US!). =Drymonia ma­ A(loplectus tetragonus (Hanst.) Hanst., Linnaea crantha (Donn.Sm.) D.N.Gibson, Phyto­ · 34: 368. 1865. Columnea tetragona (Hanst.) logia 23: 336. 1972. Kuntze, Revis. Gen. PI. 2: 472. 1891. Cran­ Alloplectus varelanus (Cuatrec.) Cuatrec., Trab. tzia tetragona (Hanst.) Fritsch, Nat. Ptlan­ Mus. Nac. Ci. Nat., Ser. Bot. 33; 120. zenfam. 4(3b): 168. 1894. TYPE: Fig. 49 in Crantzia vare/ana Cuatrec., Trab. Mus. Hanstein, Linnaea 26: 208, 215, pI. 2. 1854. Nac. Ci. Nat., Ser. Bot. 29: 13. 1935. TYPE: =Glossoloma tetragonum Hanst., Linnaea Colombia-Tolima, Ibague, 2600 m, 12 26: 208-209. 1854. May 1932, J. Cuatrecasas 2047 (holotype: A(loplectus tetragonus (Oerst.) Hanst. The bas- MA, photo: US!). =Glossoloma ichthyod­ ionym of this name has been incorrectly at­ erma (Hanst.) J.L.Clark. tributed to Oersted in the literature (Wiehler Alloplectus ventricosus Donn.Sm., Bot. Gaz. 25: 1973, Skog 1999) and on herbarium speci­ 154. 1898. =Drymonia coriacea (Oerst. ex mens, e.g., Alloplectus tetragonus (Oerst.) Hanst.) Wiehler, Phytologia 27: 325. 1973. Hanst.; Glossoloma tetragonum Oerst.; or Alloplectus villosus Hanst. in Martius, Fl. Brasil. Nematanthus tetragonus Oerst. The figure 8: 407, pI. 66, fig. 2. 1864. Hypocyrta viZ­ and description are attributed by Hanstein losa Klotzsch in Martius, Fl. Brasil. 8: 407. (1854) to Oersted as "Nematanthus tetra­ 1864, nom. nud. pro syn. Crantzia villosa gonus Oerst." Hanstein included a cited il­ (Hanst.) Fritsch in Engler & Prantl, Nat. lustration from the yet unpUblished manu- Pflanzenfam.4(3b): 168. 1894. =Nematan­ · script of Oersted in the species description. thus villosus (Hanst.) Wiehler, Phytologia ! Nevertheless, the basionym was first vali. 27: 326. 1973. : dated by Hanstein. =Glossoloma tetragon­ Alloplectus vinaceus (C.V.Morton) D.N.Gibson, i urn Hanst., Linnaea 26: 208-209. 1854. Phytologia 23: 334. 1972. Pentadenia vi­ A(loplectus teuscheri (Raymond) Wiehler, Bail- nacea (C.V.Morton) Wiehler, Phytologia 27: · eya 18: 136. 1971 [1972]. Hypocyrta teus­ 315. 1973.=Columnea vinacea C.V. cheri Raymond, Bot. Not. 113: 260. 1960. Morton, Contr. U.s. Natl. Herb. 29: 38. TYPE: Cultivated collection grown from cut­ 1944. tings at the Montreal Botanical Garden. Alloplectus viridis (C.V.Morton) D.N. Gibson, I Wild Origin: Ecuador: halfway between Phytologia 23: 335. 1972. Columnea viridis i Guayaquil and Cuenca, near I.N.C.A. camp, C.V.Morton, Contr. U.S. Natl. Herb. 29: 38. 1060 m, M. Raymond s.n. (holotype; 1944. =Columnea purpurata Hanst., Lin­ MTJB !). =Drymonia teuscheri (Raymond) naea 34: 386. 1865. I J.L.Clark, comb. nov. Alloplectus vittatus Linden & Andre, TIL Hort. A(loplectus tigrinus (Karst.) Hanst-, Linnaea 34: 17: 72, pI. 13. 1870. Crantzia vittata (Lin­ : 366. 1865. Heintzia tigrina Karst., Ausw. den & Andre) Rechinger, Oersterr. Bot. Gew. Venez. 34-36, pI. 11. 1848. Colum­ Zeit. 49: 182. 1899. =Corytoplectus specio­ nea tigrina (Karst.) Kuntze, Revis. Gen. PI. sus (Poepp.) Wiehler, Phytologia 27: 313. 2; 472. 1891. Crantzia tigrina (Karst.) 1973. : 208 SELBYANA Volume 25(2) 2f05

Alloplectus zamorensis Linden & Andre, Ill. Clark, J.L. and L.E. Skog. 2002. Novae Gesneriadeae Hort. 19: 352, pI. 110. 1872. TYPE: Plate Neotropicarum VIII: Alloplectus grandicaly*- a 110. There are no cited specimens, but the new species from Colombia and Ecuador. No~on 12(2): 173-178. I locality is described as being near Zamora Clark, J.L., L.E. Skog, and I. Salinas. 2005. Nqvae of New Grenada (Panama, Ecuador, Colom­ Gesneriaceae Neotropicarum XIV: four new $pe­ bia, and Venezuela). There is a Zamora in cies of Alloplectus from South America. No~on Ecuador where I have done extensive field­ 15(1): 70-79. i work since 1994, but there are no docu­ Clark, J.L. and E.A. Zimmer. 2003. A preliminary phy­ mented herbarium collections of this spe­ logeny of Alloplectus (Gesneriaceae): imp1icat~ons cies from this region of Ecuador. The plate for the evolution of flower resupination. Syst. ~ot. and description accurately describe a spe­ 28(2): 365-375. i cies more commonly collected from Vene­ Clark, J.L., P.S. Herendeen, L.E. Skog, and E.A ziim­ mer. In press. Phylogenetic relationships andige­ zuela and Colombia. =Corytoplectus con­ neric boundaries in the Episcieae (Gesneriache) gestus (Linden ex Hanst.) Wiehler, Phy­ inferred from nuclear, chloroplast, and morPho- tologia 27: 313. 1973. logical data. Taxon 55. i Don, G. 1837 [1838]. Gesneriaceae. Pp. 643-66$ in ACKNOWLEDGMENTS J.G. and F. Rivington et aI., eds. A General -fIis­ tory of the Dichlamydeous Plants [A General Sys­ I gratefully acknowledge the many years of tem of Gardening and Botany]. Rivington & ro., support and advice provided by Laurence E. London. I Skog of the Smithsonian Institution and Patrick ESRI. 2000. ArcView GIS, version 3.2a. Enviromhen­ S. Herendeen of George Washington University. tal Systems Research Institute; http://www.esri.9om, I also thank John Boggan, Dan Nicolson, Mi­ Redlands, Calif. I chael Nee, Christian Feuillet, and Harold Rob­ Feuillet, C. and L.E. Skog. 1990. Proposal to e~end the type citation of 7860 Alloplectus Mart., nrm. inson for their guidance in using botanical no­ cons. (Gesneriaceae: Gesnerioideae). Taxon 139: menclature and Eric H. Roalson and James F. 133-134. ! Smith for their valuable comments on the man­ Fritsch, K. 1893-1894. Gesneriaceae. Pp. 133-i144 uscript. Support for this project was provided by (1893) and pp. 145-185 (1894) in K. Prantl,1 ed. the Elvin McDonald Research Endowment Fund Die Nat. Pflanzenfam. Engelmann, Leipzig, ui­ family Gesneriaceae. Gesneriana 1(1): 1-4. ana. Acta Bot. Need. 7 [October]: 291-444. ~e­ Chautems, A 1988. Revision taxonomique et possi­ printed from Bot. Mus. Herb. Rijks Univ. Utrecht bilites d'hybridations de Nematanthus Schrader 146 [June]: 291-444. I (Gesneriaceae), genre endemique de la foret coti­ Mabberley, D.J. 1981. Edward Nathaniel Bancr~ft's ere bresilienne. Dissertationes Botanicae. Disser­ obscure botanical pUblications and his fat~er's tationes Botanicae (Berlin: J. Cramer.) 112: 1- plant names. Taxon 30(1): 7-17. ! 226. Martius, C.RP. 1829. Gesneriaceae. Pp. 27-73,! pI. CLARK: ALLOPLECTUS MONOGRAPH 209

212-226 in Nova Genera et Species Plantarum. Smith, J.E 1994. Systematics of Columnea section Impensis Auctoris, Munich, Germany. Pentadenia and section Stygnanthe (Gesneri­ ---. 1832. Gesneriaceae, nova genera et species aceae). Syst. Bot. Monogr. 44: 1-89. plantarum. Pp. 191-198 in Index Plantarum AI­ ---. 2000. Phylogenetic resolution within the tribe iarumque Rerum His Tribus Voluminibus Conten­ Episcieae (Gesneriaceae): congruence of ITS and tarum. Impensis Auctoris, Munich, Germany. ndhF sequences from parsimony and maximum­ McVaugh, R. 2000. Gesneriaceae. Pp. 253-256 in Bo­ likelihood analyses. Amer. 1. Bot. 87(6): 883-897. tanical Results of the Sesse & Mocifio Expedition Smith, J.E and K.J. Sytsma. 1994a. Molecules and ([787-J 803). VII. A Guide to Relevant Scientific morphology: congruence of data in Colwnnea Names of Plants. Hunt Institute for Botanical Doc­ (Gesneriaceae). Plant Syst. & Evo!. 193(1-4): 37- umentation, Carnegie Mellon University, Pitts­ 52. burgh, Penn. ---. J 994b. Evolution in the Andean epiphytic ge­ Moore, H.E. 1955. Drymonia macrophylla. Baileya nus Columnea (Gesneriaceae): II. Chloroplast 3(3): 109-112, fig. 43. DNA restriction site variation. Syst. Bot. 19(2): Morton, C.Y. 1938. Gesneriaceae. Pp. 1137-1187 in 317-336. P.C. Standley, ed. Flora of Costa Rica. Publ. Field ---. 1994c. Evolution in the Andean epiphytic ge­ Mus. Nat. Hist., Bot. Ser. 18. nus Columnea (Gesneriaceae): I. Morphological ---. 1963. A revision of Trichantha (Gesneri­ variation. Syst. Bot. 19(2): 220-235. aceae). Contr. U.S. Natl. Herb 38: 1-27. Smith, J.F. and C.L. Carroll. 1997. A cladistic analysis Oersted, A.S. 1858. Central americas Gesneraceer, et of the tribe Episcieae (Gesneriaceae) based on systematisk, plantegeographisk bidrag til Centra­ ndhF sequences: origin of morphological charac­ lamerikas Flora. Pp. 1-78, pI. 1-11, preprint of ters. Syst. Bot. 22(4): 713-724. 77-152, pI. 1-11. 1861. Kongel. Danske Vidensk. Smith, I.E, J.C. Wolfram, K.D. Brown, C.L. Carroll, and D.S. Denton. 1997. Tribal relationships in the Selsk. Skt., Naturvidensk. Math. Afd. Ser. 5. Gesneriaceae: evidence from DNA sequences of ---. 1861. Central americas Gesneraceer, et syste­ the chloroplast gene ndhF. Ann. Missouri Bot. matisk, pJantegeographisk Bidrag til Centralamer­ Gard. 84(1): 50-66. ikas Flora. Kongel. Danske Vidensk. Selsk. Skr., Stafleu, EA. 1956. Nomenclatural conservation in the Naturvidensk. Math. Afd. Ser. 5(5): 77-152, pJ. phanerogams. Taxon 5(5): 85-95. I-II. Stafleu, EA. and R.S. Cowan. 1981. Taxonomic Lit­ Rafinesque, C.S. 1814. Vireya sanguinolenta. Bot. erature. Bohn, Scheltema & Holkema dr. W. Junk Specchio Sci. 6: 191-195. b.v., Publishers, The Hague. Scopoli, J.A. 1777. [Gesneriaceae]. Pp. 168, 171, 173 Wiehler, H. 1971 [1972]. Name changes for some in Wolfgangum GerIe. Introductio: Historiam Na­ American Gesneriaceae in cultivation. Baileya turalem. Bibliopolam, Prague, Czech Republic. 18(4): 133-138. Skog, L.E. 1978 [1979]. Flora of Panama, Family 175. ---. 1973. 100 transfers from Alloplectus and Col­ Gesneriaceae. Ann. Missouri Bot. Gard. 65(3): wnnea (Gesneriaceae). Phytologia 27(5): 309- 783-996. 329. ---. 1999. Gesneriaceae. Pp. 492-507 in P.M. Jor­ ---. 1975. Three new species of Trichantha from gensen and Susana Le6n-Yanez, eds. Catalogue of Ecuador and Peru (Gcsneriaceae). Selbyana 1(1): the Vascular Plants of Ecuador. Monographs in 36-43. Systematic Botany from the Missouri Botanical ---. 1977. New genera and species of Gesneri­ Garden Vol. 75, St. Louis, Missouri. aceae from the neotropics. Selbyana 2(1): 67-132. Skog, L.E. and L.P. Kvist. 1997. The Gesneriaceae of ---. 1981. New species and name changes in neo­ Ecuador. Pp. 13-23 in H. Balslev, ed. Estudios tropical Gesneriaceae. Selbyana 5(3,4): 378-384. sobre Diversidad y Ecologia de Plantas, Memorias ---. 1983. A synopsis of the neotropical Gesneri­ del II Congreso Ecuadoriano de Botanica. Ponti­ aceae. Selby ana 6(1-4): 1-219. ficia Universidad Cat61ica del Ecuador, Quito, Ec­ Zimmer, E.A., R.H. Roalson, L.E. Skog, J.K. Boggan, uador. and A. Idnurm. 2002. Phylogenetic relationships Smith, J.D. 1899. Undescribed Plants from Guatemala in the Gesnerioideae (Gesneriaceae) based on and Other Central American Republics XXI. Bot. nrDNA ITS and cpDNA trnL-F and trnE-Tspacer Gaz. 27: 434-443. region sequences. Amer. J. Bot. 89(2): 296-311.